THE TAXONOMY CF THE GENUS ASTRANTHIUM (‘QOME’OSE'FAE-AS'FEREAE) Thesis for the Degree of Ph. D. ‘MICHiGéAN STAT-E UNIVERSlTY Dzi.ederik Corneiis {Dignus De Jong 39964 - -. ‘~q—. Ist This is to certify that the thesis entitled THE TAXONOMY OF THE GENUS ASTRANTHIUM (COMPOS ITAE- ASTEREAE) presented by Diederik Cornelia Dignus De Jong has been accepted towards fulfillment of the requirements for Ph. D0 degree in Botany Date 6 August 1964 0-169 Major professor A.“ ‘_ J LIBRARY Michigan State University ROOM USE NLY ROG-far“; USE GiiLY ABSTRACT THE TAXONCI‘IY 0F TIE ems ASTRANTI-m (CWCBITAE-ASTERKAE) by mederik Cornelia menus De Jong The genus gtrsnthim of the family Composites, tribe Astereae, subtribe Asterinee, consists of perennial, biennial, and annual herbs. indigenous to Mexico and the south-central and east-central United States. Ten quacies are now recognized, of which three have been newly described and one has been given a new name. Host tbxicsn species are montane and occupy cool, moist, habi- tats at elevations between 1700 and 2900 meters. me most primitive species, A. m is found on relatively dry sites in western Mexico at elevations between 800 and 1500 meters. 'me mper alti- tndinal limit or the genus 1: reached by A. Romanii at alpine eleva- tions of about 3700 meters on Gerro Potosi in the Sierra Madre wientsl. or the two species in the thited States, one, A. integ- _1'_o_l_._ig_, consists of two subspecies m1: twin-Mine in the zone of overlap. Ho species or Aetrsntlmm have been found to be metric snd‘no natural with hybrid. are imam. ' Ghraeosoue nubers are now available for all ten of Astrsnthiun. or the ten species in the genus, five are diploid, threeare tetra- ploid, me is haploid, and one is octoploid. Gin-mom studies have indicated that the basic ohro-osase nubers of gtrenthiln are _x_ - 3, h, and 5. The species with chraloso-e nubers on a base of Diederik Cornelia Dignus De Jong' J‘s-Sara themostprimitiveinthe genus, those vdthg-hare in- temediate in morphological characters, while those on 5 - 3 are the most advanced. Morphological, keryolowal, and anatomical evidence supports tin postulate that a stepdse, aneuploid reduction in basic chromosome masher has taken place in Astmthim. The reduction from isStoE-hisini'erred, but theg-htog-3drcpis supported by direct cytogenetic evidence obtained from experimental crosses . £22 thin is thought to have originated in western Mexico during the Pliocene. It became more widezpread during the Pleistocene in response to cooler climatic conditions which allowed populations to migrate along montane avenues of dispersal. Geographic isolation is thought to have been a major factor in speciation in the genus. The closest generic relatives of Lat—ranthim are mensendia and memmma. 'lhe subtribal positions or the em. traditionally placed in the subtribe Bellidinac, lstranthiun being among these, have been investigated. It is conclmd on the basis of studies of barbarian specimens that the Bellidinae' are a heterogeneous, arti- ficial group, and evidence is presented for the placement or these genera in other subtribes of the Lstereae. gtranthina has been placed in the subtribe Asterinae near Maudie and mohaetghors. Four tan previously placed in m thins are excluded from the gencsandmconsideredtobeme-bersctthegenusw the appropriate transfers have been made. m metal! 01" THE GENUS AS‘MNTHIUH (OWOSITAE-ASTW) By Diederik Cornelia Dionne De Jong A Thesis sclnitted to ’ Iticrdgan State admit: in partial fulfillment at the require-cote for the degree or 900me Department of Botany and Plant Pathology 1961; ACKNWS The present autograph could not have been completed were it not for the assistance or other minds and hands. I am.sincere1y‘graterul to the following persons: To Dr. John Ht Beanen.rcr his guidance and friendship during the course of this study. He supplied.the facilities that made possible fieldwork in the mited States and nonco and later gave invaluable aid.hy obtaining data cn.critical specinens of Astranthiun and other Astereae in European herbaria. To Dre. E. S. Beneke, J. E. cantlon, G. B. Wilson, and J. it. "right for their cosmonts and criticisms concerning the manuscript; to Dr. B. L. turner tor the use of unpub» lished chronosaae counts or several species of Astranthiun; to Drs. Harriet O. Barclay, B. B. Giannell, and B. 3. Names]: for supplying seeds or specimens; to Dr. Hladen labelin.fcr editing the Latin diagnoses and for his assistance in the library; to Mr. E. x. Longpre for new ideas and suggestions and for his assistance in the field in Woo, and to ttr. Beanie Jackson for illustrating the Mexican species. I as also grateful to the curators of the herbaria tree which speciaens were borrowed. these herbaria are listed at the beginning of the systematic treatment. The study was supported.by RationAl Science Foundation Grant 0-th while the author held a research assistantship at Riordan State university. -11- mm OF OONBN'IB um PA“ .0OOOOOOOOOOOOO...0....0.00.00...OOOOOOOOOOOOOOOOOOO ACKNOWIEDGEDEN'IB .............................................. ME OF CON'IBN'IS ............................................. LIST OF TABLES ................................................ LIST OF ILLUS‘IRA’HOIB ......................................... mmomcuon .................................................. HIS'MIIGAI. ACCOUNT ............................................ lemme AND HE'IHODS ......................................... 01101001001. INVES'HGA'HONS .................................... A. Chronosou Numbers ................................. B. Karyotype Studies .................................. 0. Discussion ......................................... CROSSING S'IUDIES .............................................. A. Interspecitic Croeeee .............................. B. Intraspecii‘ic‘croeeee .............................. WHILE!!! ................................................ A. Vasculation ........................................ l. RayFlcrets ................................. 2. Disk Flcrets ................................ B. Histolog ....fi...................................... 1e RayCa‘olh eeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeee iii a essssamsfiaEsaa sass-5e h? 52 52 iv Table of contents - w. 2. Disk Corolla ................................ 3. Pullaries .................................. WI .................................................... cremation LID BOOM! ...................................... EVOLVE!“ ..................................................... ‘0 m .00.0.0.0000...OOOCOOOOOOOIOOOOOOOOOOOOOOO... B. Evolutionary Trends ................................ cantons canteens ............................................ OHM NAHUM: ......................................... rm sum-arm. P031110! 01‘ mm: m nrsrosmce (r r! emu 01‘ m WW ...................... ....... EXCLUDED WES .............................................. 31mm W .......................................... lay te tin Qeciee ..................................... l. A. m ....................................... 2. A. ”tin.“ 0.00.00....0000......OOOOQOOOOOOOOOOOO is. A. unthece-oides .................................. 5.. 4- We“. W 5s. ;. W eehep. eiliatu 6. A- gland-e ....... s. 5. m 9. A. W. OOOOOOOOOOOOOOOOOOOOOOIOOOOOOOOOOOOOO 73 78 82 87 93 §§§E§§§E§§s Table of Contents - continued. Page 10. A. m O...00......OOOOOOOOOOOOOOO00.0.00....0. W mmm cm 0.0.00.0...0......0....OOOOOOOOOOOOOOOOOOOOOOO 171 Il'ahle l. 2 . 3. S. 6. LISTOFTABIBB heiotic chruoeue ushers in ”math. ............ thesureeente of the eonatic chi-cacao.” or the tan or mum. .......................................... amia- or Astereae with tee-bundled ray acheees and four-bmdled ray cerellae ............................ timber or vascular bundles in disk acheaes of species at Astereae .......................................... Evolutionary trends in Mgr-antid- ................... Proposed disposition er the genera of tb tribe letereae, euhtribe hllidinae ........................ pi 20 1:3 “It 85 Plate 1. 2. 3. 1;. Se 6e 7. LIST W WHWS Portion of the holetype (w shay P) of Mantra. iatmgoliu (Iticth lutt. , the type species or £traathiu ................................. Pig. 1 - ll. neietic chrenoeenes er gtranthi ....... Pig. 1 - 6. biotic and aitotic cmfiguretions in gtranthim into-“gent .............................. the karyotypes o! the taxs or gtranthilsa .............. Chart shoeiu the uperinental crosses in y_tranthiu .. Pig. 1 - 9. Chronosue configurations in superimtal m. 0: 2mm. eeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeee Pig. 1 - 6. Details of the vasculation of the achenes of m— eeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeee as. 1-6. Detailsetthvascelationetdiskaehenes G: M1... “m.— eeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeee M‘ m .1 As 1% eeeeeeeeeeeeeeeeeeeeeeeeee fibit m at £0 hCW- eeeeeeeeeeeeeeeeeeeeeeeee Geographic dietrihutiu e: A. m 5. laetiricu, 5. W and A. methane-aides ................. but draining or A. W ....................... mat drawing or A. gamma Geographic astributien e: A. M! subsp. at;- M 5- engines-em seen «u- m eeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeee '11 “8' 98 10h 5E§ 1.28 viii List or Illustrations - continued. Plate 15 . 16. 17. 18. “bit druid." of Ae Rm eeeeeeeeeeeeeeeeeeeeeeeeee Geographic distribution or A. glendens, A. Beananii, Ae "than, all As Wt... eeeeeeeeeeeeeeeeeeeee 81bit m of 5' 2m eeeeeeeeeeeeeeeeeeeeeeee Eb‘.‘ W .I A. amt“ eeeeeeeeeeeeeeeeeeeeeee Page 11:8 153 156 INTRODUCTIQI Aetranthiun, a genus of the Coupositae, tribe Astereae, sub- tribe Asterinae , is indigenous to Mexico and the south-central and east-central thited States. the nnerioan monies are well represented in the major herbaria, but a disproportionately well number of col- lections is available for the Modem species. The genus has often been confused with other genera of the Astereae which it superficially resembles. Shinnera (l9h6a) wrote: "the taxonomy of the Composites has suffered greatly from the ex- cessive reliance on the nature of the pappus, or its presence or absence, as a character of generic value”. this factor has undoubt- edly been remonsible for much of the confusion in the tamamr of mmm. m the basis of the reduced or absent pappus, mains was long considered to be congeneric with the European 21133. Also, Inrsen (1933) in her monograph of flanthiun, used this character as the widow generic criterion, placing in the genus species which had only the pappus character in common, but which differed greatly not only in other floral characters, but also in vegetative characters. the present study was undertaken with the intention of attempt- ing to clarify both intragenerio and intergeneric relationships. For this purpose, various biosystenatic avenues were explored. Cin'omo- sons nmbers, karyclcgioal data, crossing studies, and anatomical studies have been eq>ecially helpful in assessing ptnrlogenetic re- lationships. 1h adiition, an effort has been made to correlate the -1- -2- possible evolution of Astranthiuin with climatic and orogenio events in the areas in which the genus occurs. All tan, except A. robustum, were studied in the field, but not all could be studied with the same intensity. Although I had at my disposal a considerably larger number of specimens than did Larson (1933), and, in addition, benefitted n-cm actual contact with the genus in the field, mam problems need further exploration. It is hoped, however, that the present monograph will contribute to a better understanding of problems of classification in the Astereae. Plate 1. Portion of the holotype (Hichaux _s_. _n_., P) of Astranthiun Wm (Michm) Nutt., the type species of Astranthiun. he- produced fran a Kodachrome slide taken by Dr. John H. Boamm at the Museum National d'mstoire Naturalle, Paris. “I. r——-——»__..-;:: w “mmmmm..e fl HISTORICAL ACCOUNT Details of the history of Astrsnthiun following its annual description by Nuttall (181:1) may be found in the nmograph of the genes by Loreen (1933). Astrsnthiun was founded by Nuttall m the basis of a single species, A. integgifoliua, Which was first described as ge_l_1__i£ inteflolia by mohaux in the flora Boreali Americana and outsidered by him to be congeneric with the English Daisy, _3_. arenni . Apparently the only extent plants of the original material upai which Richaux based the species are cmtained on a single sheet in the Barbarian of the 14me National d'mstoire Naturelle, Paris (P). I an indebted to Dr. John H. Beaman for photocopies of this specimen (see Plate 1). The Paris collection bears the binomial Elli-.3. inteflfolia on the label as well as a Latin description of the species. The last two sentences at the label read i... in nense neio floret Guberlend“, cmstituting the sole collection data. Although these data are adnittedly vague, I have been able to obtain informatim relative to the possible locality and year of collection through scrutiny of mohaux'e Joumel (in Sargent, 1889) of his North Aeoricen travels. In the Flore Boreali Aeericene, lichen: indicated _B_e_11_i_g 1M- ggi; to occur «a ripee rivulorun et in oollibus unbrosis Tennessee“. Michsux botanlsed extensively in lorth herica between the years 1785 end 1796, end in 1795 he traveled through moon... on the hth of June, 1795, he left Knoxville m a journey to Nashville, where he arrived 12 days later. Although In crossed the Cumberland Rom -5- . -5- and Guberland River, it is most likely that the name “Guberland” on the herbariun label refers to the environs of Nashville since he calls the city ”la capitals des Etablissenents de unberland". Policing his arrival in Nashville, Nichaux spent several days collecting plants and animals around the capital and it is most likely that he collected §_e_11_i_._s_ intflolia at that tine. that he collected the species in June, late in the flowering season, is borne out by the absence of the basal and lower cauline leaves and the fruiting condition of most of the beds. "In nense naio floret' undoubtedly refers merely to the math durim which the species flowers nost abundantly. Collections of 22-3.! integgolia were also made by Thales W and c. S. Rafineaque. Refinesque (1837) outsidered Drmnmd's specimens fron mes, some of which apparently had been sent hin by John Torrey, as distinct tron g. integgfolia, and he described then as g. £13.32. In addition, Refinesque described specimens from Kentucky and lbnnessee as 932.114 gating and _B_. avi- _f_l_or_a_ respectively, with several liner varieties. Velloeo (1883) described three species of 21$ tron Brazil, but his descriptims and illustrations show these species to be members of the mnantheae or Belenieae. Nuttall, in Ms Genera of North herican Plants (1818), recorded ge—llie- awoke as described w mohaux (1803) but suggested that the fi... caulesoent species ought probably to be separated“. However, when Nuttell collected the species in 1819 almg the banks of the Pottoe River (nattall's spelling) near Mt slith, Arkansas, he noted in his diary (1821) '... the churning Daisey of herica... altogether corresponding in general aspect with the European species...‘ -7- Subsequent study of _B_e;_1_J;i_._s_ integgiiolia apparently convinced Nuttall that his original opinion on the status of the species was correct and in 181:1 he segregated Astranthium fran 21.1.13. e Although Nuttall considered the genus to be distinct, botanists such as Gray (1859): Chapman (1889), and Robinson and Fernald (1908) continued to place it in synonym under §_e_ll_i£ and nearly a century elapsed before Astranthium was accepted in the first monographic treatment of the genus by Larsen (1933). 318 transferred most of the Mexican species thus far referred under §_e_1_l_i_§_ to gtranthiun and recognized seven species and four varieties. or these species, five are retained in this paper. Three of her taxa, A. nesdcanum, A. madcanum var. chihuahuense, and A. 21319;, as well as Blake's (1937) A. guatenalense are excluded from the genus for reasons discussed elsewhere. I agree with Shinners (1950) that Astranthim intang- SEE-.111“. var. rosulatum is a mere growth form of var. ciliatm and it is reduced to synomnw. I have followed Larsen in recognizing Refinesque's 22.113 ciliata as an infraspecific taxon of Agtranthiun integjiolium; it is here treated as a subspecies rather than as a variety. Other tan described by Rafinesque (1837) are considered _ to be synonymous with g. integiioliun subsp. integlfolitn. A variety of A. integgolitan, var. robustun, described by Shimers (1950) frat Trans-Pecos Texas, is here elevated to specific rank. lhe present monograph recognises ten species, three of which are described for the first time. In addition the little known Chgsanthnun monoun- 3.3.1:. has been found to be an Astrantldun, but the cmbination i- nezioanun has already been used for a dif- ferent species, and it is thus given a new name. MATERIAIB AND Hermes Bud material for the stat of chrmoease numbers and meiotic chrueosone configuratims was collected in the field dtn'ing the comer of 1960 and the spring and sunmer of 1961. The buds were placed in vials containing a freshly mixed Carney solutim of 6 parts 95 per cent etinfl alcohol, 3 parts chloroform, and 1 part glacial acetic acid. nwebudcollections obtainedin1960 andinthe wring of1961were re— frigerated at 1° 0 upon retum to East Lansing, but the buds collected in the smear of 1961 were loept refrigerated throughout the entire field trip, as outlined by De Jong and Inngpre (1963). 811st were made by the aceto-camine shear technique. Chrmosmes were drawn with the aid of a Zeiss drawing apparatus under phase-contrast at an initial magnification of about 14000 diameters. Voucknr specimens have been deposited in the Michigan State University Herbariun. For karyotype studies, root tips were collected frat plants grown in the greenhouse fraa field-collected seeds. ‘lhe root tips were pretreated for one hour in a saturated solution of paradichloro- benzene, and fixed in a Carney solution of 3 parts 95 ”r cent etlvl alcohol andlpartglacialaoetio acidforfieinutes. Theywere anhaoqmnuyworomodiniwanatoopcrcr three tofiveninutes andtransferredtcleulgenforetainingforapedodofmminutes to two hours. Boot tip squash” were made according to the standard pro- cedure. ihe chraaosones stained darkly and were well maced apart for optima observation. lit-om 3 to 12 icbogrus were made of the karyotype of each species, and an average, representative haryotype was selected -3- -9- on the basis of meastn'uents with a millimeter rule. Watsl crosses were carried out with greenhouse-grown plants. The plants were first checked for self-capstibility. Young heads were bagged and allowed to anthese. Anthesed heads were either left bagged or the heads of a plant were rubbed together at least half a dosen times and rebagged after each transfer of pollen. lh both cases, seed set was observed and this was taken as an indication that the plants were self-compatible. (n this basis, only the pistillate florets could be used for crossing purposes. Pro-anthesis disk florets were renoved frm the receptacle with fine tweezers and the heads, with only the ray florets left, were then bagged to prevent contamination. Each head contained from 15 to 35 ray florets, depending upon the species. When the stigaas were receptive, pollen was transferred by rubbing intact heads with anthesed disk florets, which had been bagged before anthesis, onto the stignas. This procedure was repeated at least half a doeen times for each head during the ensuing three of four days. Following each transfer of pollen, the "female“ heads as well as tin bade of the “nale' parent were carefully rebegged. Each cross was attempted with m plants, but the met nunber of plants utilised was not recorded. Forth studyofthevasculstionoftherayanddiskflorets, both fresh and dried plants were used. Fresh material was placed in PM for killing and fixing, washed in water, and transferred to 2.5 per cent aqmous soon for clearing. n-ied material was placed directly in this clearing solutim. Glacial acetic acid was used for terlinal clearim, after winch the florets were washed and dewdrated in an etw1 alcohol series. Whole momts were prepared by relieving cleared -10- material to a 1 per cent solution of safranin in equal parts of absolute ethyl alcohol and xylene for staining, after which the florets were transferred to xylene and mounted in balsam. Flesh and dried heads were used for histological studies. Fresh material was killed and fixed in PM and dried material was softened and cleared as above. The heads were then washed and in- filtrated and embedded according to the tertiary butyl alcohol method of Johensen (l9h0). Staining was carried out with safranin and fast green according to Northen's variation of F'oster's tannic acid-ferric chloride nethod (Johansen, 191.0). Table l. Meiotic chromosome numbers in Astrarrthiua. taxon 3 number collection or reference 5. lo an 5 De Jon; and Ipngpre (1963) A. laetificum 10 HICHOACAN: 1.5 mi W of Mil Cmbres, 29. m 2.95 A. wurascens B Turner, Human, and Rock (1961); De Jena and Ipnzpre (1963) 8 CHIAPAS: ca 5 mi E of San Cristobal de Las Casae, 280 , Turner, Ellison, and unpu .) 8 cmms: Just w or gen Cristobal de Les Oasas, % 2 humor, Ellison and an; (unp 1.; 8 + 1 CHIAPAS: ca 10 mi E or Teopisca, or 2 trapents 193 £11, Turner, Ellison, and King unpu . 8 CHIRAS89m1Wo£CuI1tan D 8 GEARS: 5.2 :1 E of loopisca, 2: £33.63 8 0mm: between his in and MB, Route 190, 93 m 113; 8 WA:O.2niWo£Ellb1-tin, Route 190, 2 £23 1116 5. mthocuaoides B Turner, Boanan, and Rock (1961); De Jon; and Ionapre (1963) 5. into line 14 Baldwin (191:1, reported as _A_. u a A Table l - Continued taxon n umber collection or reference 5. into foliu h subsp. ta 0- E9. . into oliln h teasing u u 1. u A. splendene 9 9 TENNESSEE: Davidson County, ca 20 mi 5 of Nashville, Channell 8880 omen: Pittsburz County, 2 mi N or Crowder, 25 £23 119 am: Pittsburg County, 6.5 mi E of Arpellar, 9: Joy L8}, 0mm: Hughes County, In? mi E of Pontotoc County line, Q. m 192 TEXAS: Colorado County, 8 mi NW of Eagle Lake, Turner 3259, Turner (unpub1.) W: San Patricio County, 6 mi NE of Staten, m:- @163, Turner (unpub1.) IUEVO was: between m: 30 and 31, read to Dr. Arroyo, 23 «31230 mo 1.“: between me 29 and 30, road to 11-. Arroyo, pg J02; 1239 Turner, Rama, and Rock (1961, re- ported as Aetranthit- 52.) mo m: SE slope of Cerro Potosi, a 12: 52k IUEVO m: alpine meadow, peak of Cerro Potosi, .2... 929‘ 522 IUEVO m: I side of peak, Cerro MM: 2: £03 22 NUEVO mow: alpine meadow, peak of Cerro Potosi, 2 Joy 199 E slope at Cerro Potosi, 21931.82. Table l - Continued collection or reference taxon g nuber 5. «them 3 3 A. condimentun 3 3 3 3 3 A. robustua 3 De Jon; and lawn (1963) MANGO: ca 31 mi an of Ciudad mango, 3%, turner, Ellison, and Kim unpu . JALISCO: 35 mi at of Jiquilpan, 2: £23 .1311 HICHOACANa ca 33 mi W of Horelia, % #2, Turner, mison, and unpubl.) 1410mm: at n: 10.5, Route 39 from Carapan to (h'uapan, $333163 Emma“ 15 mi S of Villa Escalente, 2-; 92:. __.1°82 HICELCAI: 5.1; mi E o! Quiroga, 9.9. mm; TEXAS: Golf Course, Alpine, "flak gage Plate 2. Fig. l - ll. biotic chromosanes of Astranthimn, x 11170. Fig. 1. A. m, a - 5 (_D_s_ gnaw. Fig. 2. A. laetii'icm, g- 10 (.D_e_ 193%. Fig. 3. _A_. pmurascens, 3- 8 (211233.31)’ Fig. 1;. A. mthooomoides, g - 8 (.12. £939.22). Fig. 5. A. M- £13 subsp. inte olium g- h (Channell 8889). Fig. 6 A. _i_n_t_e_- Egolimn subsp. ciliatum, g - h (93 £93182). Fig. 7. _A_. splendsns, 5-9(ge_ge£gl2_lfl). Fig.8. -A_.Beamanii,g-l2(2.e;m1§2). Fig. 9. 5° ortho tan 3 - 3 (93mm). Fig. 10. 5. condimentm, g- 3 (93mm). Fig. ll. .15. robustua, g - 3 (W 2.9.). CYTOINICAL INVESTIGATIWS A. Chromosome W. Chromosome numbers determined from observations of pollen mother cells are listed in Table 1 (cf. also Plate 2). Heiosis was regular in all taxa examined except in one population of Astranthiuu intggg- m (2 £93 M) from Fayetteville, Arkansas. This population was morphologically intermediate between subspp. integigoliun and ciliatum. 'menty rosettes of the population were collected in the field in the spring of 1961 and grown to maturity in the greenhouse. All plants were examined cytologically: 18 plants had 2 - 1; while two hadg- 5 (cf. Plate 3, Fig l - 6). Inthese twoplantsithe fifth chromosome sanetimes appeared as a small bivalent with normal anaphase movement, but in other instances it was found attached by slender chraaatin threads to another chromosome. Root tips of the two plants all had ten chromosomes, the extra pair evident as small clu'auosomes with subterminal centromeres. (no possible explanation is that the g - 5 plants resulted from an ansuploid increase following a series of unequal translocations. In view of tin presence in Astranthiun of the basic nuber _x_ - 5, another possibility is that they are sur- vivors of an ancestral form with; - 5. The two plants could not be distinguislnd morphologically from those with g - h. meocourrenceofthebam chromosaenmbersg-S, h, and3, suggests that a stepdse, aneuploid reduction of the basic chrmosome umber of the "9221-2. type (Babcock, 19152) has taken place in -16- Plate 3. Fig. l - 6. Meiotic and mitotic configurations in m- 21.1.“; integgonun (20; £93 13,), x 1.000. Fig. 1. Normal ohrano- some nunber, g - h, metaphase, I. Fig. 2. Diakinesis, g - 5, extra bivalent indicated by arrow. Fig. 3. ’ niakinesis, g - 5, extra chro- mosome (mow) attached by chromatin threads. Fig. 1;. Makinesis, n - 5, extra bivalent shown by arrow; hatched area lightly stained. Fig. 5 and 6. Somatic metaphase, 2g - 10, extra pair shown by arrows in Fig. 5. -19- Astranthiun. Although evidence for the drop frau 5 - s to :_: - t can only be inferred at the present tine, direct cytcgenetic evidence hr the drcpfromg-h tog-3 hasbeen obtained. is indicatedin the section on experimntal crosses, F1 hybrids obtained free a cross be- tween 5. integifcliu subsp. integifolim (g - h) and A. condinntua (a I- 3) showed me trivalent all! two bivalente at dialdnesia a! micro- sporogenesis (Plate 6, Fig. l - 5), indicating haaology betwoen the four chraaoecnes cf the one parent with the three of the other. In addition, 1"l hybrids obtained from crosses between other mmbers with ctn'cmoscue nmbers based on 5 - h and _x_ - 3 shaved a high percentage of trivalent formation. his the secmd time that evidence for ansuploid reduction has been directly observed in a genus of file Astereae. Jackson (1962) fmnd a similar case in W sect. Blepharcdon. Five species of Astranthium are diploid and the other five, all Hexican, are polyplodd. Of these polyploids, three are tetraplcid, one is haxaploid, and one is octoplodd. It may be inferred fraa the regular pairing configurations of the polyploids that they are allo- poly'ploids. All polyploids are perennials. B. Karzotm Studies. Husiwara (1959. and papers cited therein) has conducted a m i of karyotype investigations in the istereae, mbu'ibe isterinae. lhe results of his smdiee have been of considerable value in determining phylogenetic trends. 'lhe diverse array of cuteness. more in g- m-enthium pranptsd me to undertake a survey of baryotypes in all taxa in order to dstsradne whether a' not evolntion of chromosome mcrpholcgy fable 2. lbasuruaents of the somatic chrancsomes of tax: of m- tancn ohratosome length in short arm position of pairs microns length/ total centrcmere length (1100) ;. g1m 1, 2 9.0 so sulmedian 3, h 8.2 h2 submedian 5, 6 3.0 no Unwed“ 7, 8 5.0 ho suhnedian 9, 10 h.0 30 subodian a. laetificm l, 2 7.0 50 median 3, h 7.0 ’10 ltfilfldiln 5, 6 6.2 50 median 7, 8 h.9 25 subteminal 9, 10 h.3 35 submedian 11: 12 1802 33 submdian 13, 1h h.0 38 comedian 15, 16 3.1; hl suhnedian 17, 18 3.0 33 suhnedian 1.9, 20 2e3 35 Cum.“ 5. murascens l, 2 5.5 143 suhaedian . 3, h h. 3 23 subteninal S, 6 h.0 38 subedian 7, a 3e8 26 Baum 9, 10 3.8 26 subtemainal ll, 12 3.1; fl «Indian 13, 11; 3.1 26 subterminal 15’ 16 105 27 Cubans-”.1 A. W 1, 2 h.l 32 subedian 3, h 3.}: 28 subtenfinal 5, 6 3.1 26 subterminal 7, s 3.0 33 suhaedian 9, 10 2e. 5 CW ll, 12 2.1; I auhnedian 13a 115 1.9 157 aubedian 15, 16 1.1; 21 subteninal -21- Table 2 - antimled tamon chromosome length in short are position of pairs microns length/total centruere length (2100) a. into 11:- l, 2 13.8 36 submedian squp. fifi- 3, h 3.8 29 subteninal foliun 5, 6 3.3 33 suhedian 7, 8 3.0 33 submedian As m 11D 1, 2 1103 33 3mm sEsp. h 3, h 3.8 29 subterminal 5, 6 3.1 32 submedian 7, 8 2.8 32 3‘1”“ '5. glendens l, 2 11.8 118 subtledian 33 h 1‘06 28 subterminal 5, 6 3.7 28 subterminal 7, 8 3.6 27 subterminal 9, 10 3.5 28 subterminal 11, 12 3.3 33 suhnedian 13, lb. 3.1 26 subterminal 15, 16 3.1 26 subterminal 17, 18 2.8 28 subterlinal A. Denali l, 2 luh 37 submdian 3, h h.3 23 subterminal 5, 6 8.3 133 subnedian 7, 8 3e6 31 811W 9, 10 3.5 28 subterminal 11, 12 3.3 33 aubmedian 13. 11: 3.2 28 subteninal 15, 16 2.9 27 subteninal 17, 18 2.8 20 subterminal 19, 20 2.8 35 sum 21, 22 2.6 31 lib-ediln 23, 2h 2.6 31. auhnedian _A_. M l, 2 3.8 26 subterninal 3, h 3.6 22 subtenninal 5, 6 2.9 31 suhaedian A. condimentm l, 2 h.0 25 subterainal 3, h 3.8 26 subterminal 5, 6 3.3 33 subnedian -22.. Table 2 - Continued tazcn chromosome length in short arm position of pairs microns length/total centraaere length (1100) A. robustun l, 2 11.1 31 sumedian 3, h 3.8 21 subterminal 5. 3.2 27 subteminal -23. paralleled that of gross morphological and anatanical characters. Procedures have been outlined in the section on materials and methods. The measlnements of the chromesues of each tamon are listed in Table 2andideograms foralltaxaareshcwnonPlateh. Observations . l. manthium m (22 - 10). misspecies is diploidonabaseofg-Sandhasthelargest chromosomes of am mecies in the genus. Gin-omosome size ranges fran 14.0 - 9.0 microns. 0f the five pairs, four hsve suhnedian centromeres, whereas the largest pair has median centraaeres. 'Ihe mnallest pair bears satellites. 2. Asganthium getificm (2g - 20). The karyotype of this tetraploid species is characterised by a predauinance of chruaosomes with suhaedian centromeres. he pairs have median centromeres and only one pair has subteninal centromeres. The second smallest pair has satellites. Mal chromosome length ranges fru 2.3 - 7.0 microns. 3. Ammthiu pmurascens (2g - l6). misspeciesistetraploidmabaseofx-h. atheight pairs of chrcaecscmes in the comlement, three pairs have subaecflmn centrueres, the mining pairs have subterminal centruneres. Mal chrmosae length ranges fro 1.5 microns to 5.5 microns. The unall- est pair has satellites. 1;. main- menthccaoides (25 - 16). in this tetraploid species, five pairs have sulnedian centrueres, ~28- Plate h. The karyotypes of the tan of istranthimn. The cin'omosomes are arranged according to decreasing total length. The species se- quence corresponds to that of the systematic treatment. .25. n K {I '8 is A.XYLOPODUM ), n H H I! u it n if u A'LAET'FICUM “ N n I: 8) n n if A.PURPURASCENS H II n n u u u n A-XANTHOCOMOIDEs " ‘i '| ‘. A.INTEGRIFOLIUM subsp. INTEGRIFOLIUM “ n u “ A.INTEGRIFOLIUM subsp. CILIATUM (I )K‘ u 0 It u m: u K0 37 H u n u n n u u u n u “ fl ‘8 H A.CONDIMENTUM u n “ -26— while three pairs have subtemninal centromeres. is in 5° pmurascens, the nallest pair has satellites. chrcucecm siae ranges free 1.). to he]. Mime. 5. Astranthimn integigcliln subsp. intemgoliu (22 - 8). me majority of the preparations which I made of the root tips showed the largest pair of chromosomes with extremely small satellites. In certain instances, however, these were not visible and must have been obscured by the short arm of the chromosomes. Chromosome size ranges from 3.0 to h.8 microns. 'L'aree pairs have submedian centro- meres, while only one pair has subteminal centromeres. A 6. ggmthimn integggolim subsp. ciliatum (25 - 8). 'me somatic chrmoscmes of subsp. ciliatum areislightly maller than those of subsp. inteflcliun. ibis is also reflected in the size of the pollen grains of the two taxa. In subsp. integEgoliun, pollen grain size ranges fran 16.0 to 20.0 microns, with an average of 17.5 microns; subsp. ciliatua has pollen grains with a siae range of 15.0 to 18.0 microns, averaging 16.5 microns. me largest chromo- some pair in this subspecies also has small satellites. 7. menu Elendens (2g - 18). weepeciesisbnploidonabaseofg- 3. thDninepairs in the caplement, seven pairs have subteninal centrmaeres, the re- maining pairs sut-edian centraaeres. No pairs of satellited chrano- somes are present. Chromosome size ranges frat 2.8 to 17.8 microns. a. matting—“mm (22'2‘0' . ibis octoploid species has elm-omcsomes which range from 2.6 to ~27- h.h microns in length. Five pairs have subterminal centraasres, where- as uven pairs have subnedian ones. As in 5. splendene, the knryotype of A. Bemnii has two pairs of satellited chromosome. 9. Aetranthiun orthogodun (2g; - 6). g. orthopodum is diploid with 6 saaatic dlranosones, four of which have subterninal oentronsres. 'lhe sunset pair has subnedian centrmsres and is 2.9 microns long. he largest pair, which bears satellites, is 3.8 microns in length. 10. Aetranthiun cmdinsntun (2! - 6). '11: six chronoeones of this diploid species range from 3.3 to 11.0 microns. As in A. crihgpodun, the smallest pair Ins subundian oentraurss, the remaining pairs have subteruinal cmtronres. me largest pair consists d‘ satellited chronceonee. 11. Astranthiun robnsmn (22 - 6). As in 5. crmopodm and A. condimenmn, the largest molasses pair has satellites, but the centromres are mbmdian rather than mbterninal. ‘me smaller pairs have subterninal centromres. Chro- nosw length ranges from 3.2 to 11.1 Iicrons. C. Diecuegeion. Chromosone number, sine, positim of the centre-ere, and pres- ume of secondary constrictions have woven useful as hryotype characters in Aetranéig. 'me chromou couplenente of A. El};- Egtg (2g - 10) and 5. lastincua (2g - 20) contain the largest chro- mosome of any species in the genus. In addition, their karyctypee are largely sy-etrical, i.e., fliers is a pepmdu'anoe of cln‘onoecnee -28.. with median centronnres and submedian cents-owes, while chromosomes with subterminal centronnres are absent or few. his condition is generally considered to be primitive (cf. Stabbim, 1950) an! is in agreeeent with the cytogenetic evidence which suggests that the basic chromosome mmbers 5 . h and 5 - 3 lave been derived throng: aneuploid loss. The species thich have chromosome nuabere on a base of 5 - h, and which are intermediate in morpholouoal characters, also have intermediate ltaryotypee. 'lhe species wifli 5 - 3, which are morpho- logically specialized, have karyctypee which are aemtzical with a predominance of subtsrminal cent-cures. these species also have smaller chromosome than the Irimitive species. ‘lhe ltaryotypes d 5. m an! 5. lastii’icum show mm simi- larities in chranoson sire and position of centrolnree. here in a sharp break in total chromosome length betteen the third and fourth pair in 5. m and in shcrt-arm length between the third and fourth pair of 5. 1aetlficum. In cmtrast, the soentic chromosome of all other tars, when arranged according to decreasing length, show a graded series. Astranthium Module and 5. laetificum are also more sharply differentiated from the rest of the genus by gross mor- phological characters than are am other species. It is perhaps also significant in this respect that attempts to reduce lubride betaveen 5. lastificum and species with 5 - 1: mad 5 I 3 (especially 5. m- mwifi: 5- 8, 5. glendenewith 5 - 9, ad 5. Beunii withg - 12) failed, while hybrids could be synthesized between species with 5 - I; am 5 - 3. ' Species which are related npholcgcally and mu have the same chromoeme number often have secondary constriction a: the cue -29.. chraloeome pairs. This is especially strihng in 5. pmuraecens and 5. xanthocanoides in which the smallest pair bears satellites, and in 5. orthopodun, 5. cmdimntmn, and 5. robustum, in which the largest pair has secondary constrictions. Astranthiun gglendens and 5. Bamanii have satellites on the second and fourth chromosome pairs. me hryotypic data are especially important in connection with these two species in supporting the assunption that their basic chromosome umber 1. 5 - 3 (of. also discussion under 5. gionoans . Although 5'9has been shownby Solbrigfig. (196M tobe themost colmon basic number in the Astereae, the occurrence of 5 - 12 in 5. Bsamanii makes it appear unlikely that 5 - 9 is a diploid nunber in the nor- phologically very similar 5. glendens. It is more liloely that 5. glendens is a hexaploid on a base of 5 - 3, and this hypothesis is supported by its advanced heryotype morphology as conquered to that of other species or Astranthiun. lilo independent, opposing trends in chromosome evolution have occurred in 5stranthium, and both trends seem to provide definite selective advantages. 0: the one hand, there has been a reduction of the basic chromosome number through anewloid loss, and on the other, an increase in chromosome number through polyploicw. The two species with the basic umber 5 - 5 are primitive perennials, one is diploid, the other tetraploid. Two polyploid species with 5 ' h are perennial, as are a Maploid and an octoploid species on the base of 5 - 3. Since these species are primarily montane, it is possible that both the permial habit and polyploiw may have a selective advantage for plants occupying cool, moist habitats in the mountains. It is perhaps significant in this reQect that the highest polyploid, 5. Beamanii, -3)- occurs at higher elevations than any other species or the genus. A correlation of the perennial habit and polyploiw with high elevations and latitudes is well known (Gustatseon, 191:8). me reasons for this phenomenon are not yet entirely clear, however. In. contrast to the occurrence of polyploids in montane areas, two of the more advanced, diploid species, 5. integgolium and 5. robustum, are annuals and are found in warmer, drier habitats at lower elevations. No other advanced species, 5. ortgom'n and 5. 505151.- ”, occur in mountainous areas, but their habitats may be special- ized in acne otter way, as is suggested by their restricted geographi- cal distribution. Blatively minor climatic changes may have had significant effects on the habitats which the species with low manbers occmy. The ability to evolve adaptations to new enviromnental conditions would be important to their survival. We would seem to be especially important with respect to the supposed Pleistocene evolution or the genus (see under Origin). The rapid expression of genes or gene combinhtions might be facilitated by the low chromosome nubers. ‘lhe annual habit of these species, which would necessitate segregation 5 and recmbinstion every year, no also have been important in in- creasing their potential for rapid evolutionary change . Solbrig 35 55. (1961;) have found that genera of the istereae with low chro- mosome mothers are especially trequent in arid areas or the Southwest and Hence. they also believe that these low numbers W represent adaptations to specialised habitats. magical and plvsiologioal changes in the species of methim may have been accompanied by and possibly, but not -31.. necessarily, facilitated by changes in the karyotype. Progressive changes from symetrical to asymtrical karyotypes and from large to median-sized to small chromosomes probably resulted from trans- locations and inversions. Perhaps it is the new gene combinations which result Iran these chromosomal alterations, rather than actual morphological changes in the karyotype itself , which have a selective advantage. Ihe alteration of the karyotype may thus reflect only the fact that a greater number of translocations and inversions have occurred in tame with an ametrical karyotype than in those in which it is symetrical. CROSSING STUDIES Ebzperimental crosses were attempted using greenhouse-grown plants of nine species of Etranthium. The crosses attempted are shown on Plate 5. Solid lines indicate successful crosses, broken lines un- successful crosses. Crossing procedures are outlined in the section on materials and methods. The Inrbrid nature of the F1 plants obtained was verified using the following criteria: (1) Their morphology was carefully compared with that of tin parental species. 1h most instances, the mbrids were intermediate between the parents in quantitative characters, while qualitative characters such as ray color were found to be dom- inant. (2) The somatic chromosome number of each hybrid was checloed while the plants were still in the vegetative stage. This was the most conclusive evidence for twbrids obtained from crosses between heteroploid parents. (3) Meiotic irregularities may be an indication of mbridity in lwbrids whose parents show regular meiotic con- figurations . (h) The percentage pollen stainability of each lybrid was always considerably lower than that of either parent. (I: the basis on! these criteria, there was never am doubt about the Ivbrid nature of the 1'1 plants. 1!» parents which were used in the successful crosses and for which percentages of pollen stainability are known are listed in tabular form below. Plate 5. Chart showing the experimental crosses carried out in green- house cultures of Astranthium. Solid lines indicate successful crosses, broken lines unsuccessful crosses. The haploid chromosome nunber of each taxon is indicated. PURPURABCENS LAETIFICUH CONDIMENTU" INTEGRIFOLIU ORTHOPODUM subsp. INTEGRIFO LIUI SPLENDENS - 35- species collection 5 pollen stainability A. Beamanii De Jo 66 99.8 '5. c ntmn 5' 3 ;E 90.0 5. or c mu 5% 9M6 _. Eggpurascens 5' 3a_ng 96.8 t BEE—miss; Es”... 35-: is 2 mns E M 95el‘ Observations on meiosis in the lubrids were made on pollen mother cells stained with aceto-carmine and observed on slides made according to the routine squash technique . Tl'm percentages of pollen stain- ability are based on a total of 1500 to 2000 grains frat one to several heads frat plants in culture. Pollen grains were stained with cotton- blue in lactophenol for a minimum of twelve hours. The percentages are based upon the number of grains which stained dark blue. The Inbridisatione are grouped into two categories: (1) crosses between morphologically distinct, closely or more distantly related heteroploid or hanoploid tame, and (2) intraspecific crosses. Time was not available for crosses beyond the F1 generation. The crosses in which Fl lybrids were obtained are discussed below; the pollen re- ceiving parent is listed first. A. Interspecific Crosses 1. 5. integ‘golim subsp. intezgolim (_ - h) 1 5. condimentma (9, - 3). ' 'lhe plants of 5. intemgun subm. integrgolim were groin from seed of plants supplied by 11‘. R. B. Channell (Ohmell 8880). -36- Sven ecstatic chranoeomes were present in the root tips of the two P1 plants obtained (Plate 6, Fig 7). In meiosis, 50 cells showed one trivalent and two bivalente. At diakinesie, this trivalent was present as a ring or a chain (Plate 6, Fig l - 5). mmetines a single univalent and a loosely associated bivalent were found, but such con- figurations were extremely rare . (he trivalent and four univalcnts were observed in only two cells. Pollen stainability was a mere 2.9 per cent. This cross provided tin most conclusive evidence for aneuploid reduction in gtranthime. 2. 5. ortmodun (5 - 3) X5. condimentum (5 - 3). 1h 80 cells examined, only a few showed two univalents at dia- ldnesis, the remaining cells did not show any meiotic irregularities. Altimugh normal pollen tetrads were formed, pollen stainability was only 51;.0 per cent. In view of tie regular meiosis observed, this relatively low percentage is notewortly. Krucloeberg (1961) recently made similar observations on artificial lubrids between closely re- lated species of _S_i_le__ne_. It suggested that the reduced fertility of hoquloid lybrids was due to "...genic ...or minute eegnentsl differ- ences, below tin threshold of detectial, in the chrauosmes". The evidence suggests that this may also be the case in this particular instance. 3. 5. pguraecens (5 - 8) x 5. condimentu (5 - 3). Th plants of 5. 232mm used in this cross were from collectiongingm. The root tip chromosomes of the three 11 plants obtained were 11 in number (Plate 6, Fig 8). As expected, meiosis was extremely irregular, with a dngle trivalent, one bivalent, Plate 6. Fig. l - 9. Chromosome configurations in emerimental tw- brids of istranthimn. Fig. 1 - 5. nialdnesis in tybrids of 5. 554- rifolium subsp. intemnum (_ - h) x 5. gm, (5 - 3). Fig. 1. Two bivalents and a circular trivalent. Fig. 2 - h. M bivalents and a trivalent-chain. Fig. 5. No bivalents, one loosely associated, and a trivalent-chain. Fig. 6. Somatic ohrusoscmes of the lubrid 5. Boamanii (_ - 12) I5. fleadene (5 - 9), showing 21 chromoeases. Fig. 7. mastic chraaosaaee of the lubrid 5. intemgolim subsp. integi_folium (5 - h). I 5. oondimentmnvglr- 3), showing seven chr00- smee. Pig. 8. Smeatio chromosomes of the lubrid 5. pmurascens (5 - 8) I 5. condimentm (5 - 3), showing 11 chraoecmee. Pig. 9. elastic chraeosaaes ofthelubrid5. mg- 3) X5. pu_r- pursscens (5 - 8), showing 11 chromosaes. ~38- .- 39- and six univalents observed in most instances. In other cells I found one trivalent, two bivalents, and four univalents, or three bivalents and five univalents. No stained pollen grains were found. h. 5. EM“. (5'- 3) X5. pmurasoens (5- 8). The plants of 5. gmurascens used in this cross were from collection 2. 5125 5535. The somatic cells of the three F1 plants clearly showed 11 chromosomes (Plate 6, Fig 9). is expected in off- spring fran parents with widely different chromosome numbers, meiosis was highly irregular and meiotic configmtions were sanetimes diffi- cult to interpret because of extreme clumping of the chromosomes. In 25 cells examined, 18 contained a single trivalent, one bi- valent, and six univalents, or a trivalent, two bivalents, and four univalents, or a trivalent and eight univalents. In other cells, the following configurations were observed: four univalents, one bivalent, and a clump of five chromosomes; two bivalents and seven univalents; a clump of five chromosanss and six univalents; one bivalent and nine univalents; rarely, ll univalents. No stained pollen grains were ob- served in the hybrids. The reciprocal cross was successful, but the single F1 plant did not reach maturity. 5. 5. glendens (_ - 9) X 5. pmurascens (_n_ - 8). Plants of 5. pguraecens were free collection _D_e_ m 512. The following meiotic configurations were observed: one trivalent and seven bivalents; me trivalent, six bivalents, and two univelente; one trivalent, five bivalents , and four univalents; six bivalents and five univalents. In thee uiotic figures, one trivalent and seven bi- valents were most frequently seen. Pollen stainability was 65.0 per cent. -50.. 6. 5. mamanii (5' 12) X5. glendens (5- 9). Five 11 plants were obtained in this cross. The root tip chro- mosomes are shown on Plate 6, Fig 6. Attempts to analyze microsporo- genesis failed because of the extreme clumping of the chromoemuues, and observations are therefore limited. In addition to the chromosome clump, fran one to five univalents were seen. Lagging chromosomes could be seen in first and second anapbsse, and micronuclei formation in tin tetrads was frequent. Pollen stainability was 58.5 per cent. 7. 5. 21endens(5 - 9) :5. ort_h_ogodun (_ - 3). The two 171 plants obtained in this cross showed 12 somatic chrasosomes. The lybrids are approximately one year old and are still in the vegetative stage, consequently, it has not yet been possible to analyze meiosis. B. lhtraspecific Grosses Early in this stuw, an attenpt was made to ascertain whether the three widely separated populations of flmthiumu gym-sesame are specifically distinct. The following intraspecific crosses were carried out: f-ale parent male parent tubs-id pollm'u by collecticun by collection stainability S 2mm (m) leaaflhncl) 1.5 55 551‘ 152 (Oar-ca) i 95 m 5555 (Chiapas) 8.2 2. 51315155 (Chiapas) I 25.12532 (Oaxaca) N 1.3 2'. as 1.211 (maniac) x 2 £53 all; (Chilpas) 3.1 -hl- The three populations or A. gmurasoens are represented by collection ntsnbers 2'. go}; $231, 32, and g respectively. No differences could be observed in chromosome number and karyotypes in these collections. Taxonomic studies have indicated that there is an overlap of morphological characters, consequently, A. pmurascens is here treated as a single, it somewhat polymorphic species. A stuck of microsporogenesis of each IHybrid" rarely showed an irregularities. The extremely low percentages of pollen stainability are therefore surprising. Although the three populations have not yet acMeved morphological distinctness, genetic barriers appear to be present, in part possibly as a result of geographical isolation. The regular meiosis in the hybrids and the similarity in the loaryotypes of the three populations would suggest that mall, genie or seguental difierenoes in the chrmosmes are responsible for this reduced lubrid fertility. MORAL mm The anatomical studies reported here involve both the study or whole-mount material for the purpose of caparing floral and achenial nodes or vasculation, and observations on the histology of the capi- tula or Astranthium and species or other genera in the Astereae. A. Vasculaticn 1. £51 1.1.2223“ nae ray corollas oi' £tranthium are ligulate and consist of a short, sparsely pubescent tube and a flattened, ex- panded lamina with the apex cosmonly 3-toothed. The venetian oi” the lamina typically shows tour vascular bumlles which originate fro the vascular plexus below tin apex of the ray achene. 'nasse bundles traverse thelengthotthelaminaandconvergeattheapem. m corollas of some ray florets may show five, or less calmly six or even seven veins, but the tour-bundled condition is by far the com- monest in the genus. his type of corolla vaccination is the ”Actor- WPO" 01' 50011 (191))- At the base or each ray scheme, a circle vascular bundle enters true the receptacle. Upon entering, this bundle divides to tom two lateralbmdleswhiehtraversethe lengthotthe aobne. A single ovuletrece‘naysuetinesbeseenbranohingottthebasallmotot vascular tissue, but this truce extends upwlrd.1br'only a short dis- tance. melateralachenialbmdlesnaycmsistota singlerowot tracheary elements or may contain several, mostly Wt thinner, tiles which co-only increase in member and consequently in thickness .hz- Table 3. anecies of Asterese with two-bundled ray achenes and four- bundled ray corollas. species source of material (MS) Achaetoeron m A. fleeus ELLE. Egremis 2. glwstris Brac ace tree a Chaotggpga 9229.922. nichaetoghora cm atria lbwnsendie lexicana 2- man a- m .0.» an; a My: .1922 92 £23 a Lamas 2E m 2931 Botanical Garden, Coimbra , Portugal (14$ 166628) 2222.8. $2.6. ma £2.22 £21.95}: £22 m l§§1 m 1.4322 Pgson §t_ Amatm 29E 4,1,- toward the apical plexus. 1h cross-section, the two nerves of each aclnne may be seen as small areas or sclerixied cells, each directly opposite and external to a bundle. At the plexus, the two bundlu branch, each toning two corolla bundles and one stylar bundle. A1- thowhthe achenialbmdlesappeartoheireetromoneanotheratthe plexus, they are occasionallyintercomected. A third, thinner bmdle with a single file of traclnary elements has been observed in sane ray schemes of A. ortmodun, and this bundle branches independently into a corolla bundle and a stylar bundle. mo vascular bundles are found in the ray schemes of all species or Astranthiu except 5.- glggodua and A. lastirictn. whereas all other species have two-normed schemes, the schemes or 5. 1122232". and _A.. laetifloun are prominently ribbed and the mode of vasculation in these species seems to be correlated with the position of the ribs. The rq schemes of A. M are two- or three-ribbed, and have achenial tacos which may be provicbd with one or two conspicuous longitudinal nerves. 1!: cross-section, these ribs and nerves appear as areas of bavily sclerii‘ied cells. lbur vascular bmdles, two lateral and two (lord-ventral, branch of: a single trace which enters the base of the scheme from the receptacle. 1h two-ribbed achenes which have a single nerve on each face, each lateral bmdle is dir- ectly opposite a rib and each dord-ventral bundle is directly oppositeanerve. tuntbrqacbnesarethree-ribbed, onslateral bulileisoppodtemerib, idletheotheris aboutddwayand alternate with tin two remaining ribs. me dorsi-ventral bundles are again opposite a nerve, if present. lh A. laetinclle, the ray echoes are two- or three-ribbed. Two- Plate 7. Fig. 1 - 6. Details or the vasculation of the achenes or Astranthium, viewed laterally. Corolla bundles, 3; style bundles, g; anther bundles unlabeled. Fig. 1. Ray achene of A. orthoflm, X 80 (313M210. Fig. 2. Disk achene of A. gleam, x 80 (2.1% _l_9_2_). Fig. 3. Disk achene of _A_. laetificmn, 1 ho (21mg). Fig. 1;. Disk achene of A. xanthocomoides, X 80 (De Jong 1195). Fig. 5. Disk achene of A. orthopodun, x 80 (2,13%). Fig. 6. Disk achene of A. pmurascens, X 80 (p; Jogg M). -57- ribbed achenes have two vascular bundles, each bundle directly situ- ated opposite a rib. Three-ribbed acbnes have three vascular bundles, eachbundle sgainqaposite arib. Ihlike tb actuatedbm the ray achenee of A. laetificu have nerveless faces. the mode of vasculation of the ray florets was investigated in detail only in gtranthiu, but limited observations were also made of cross-sections of heads of species of other genera in the Astereae. 1b "Aster-type' of corolla vasculation (Koch, 193)) was also observed in tin species which are listed in'l‘able 3. the ray schemes of these species also appeared to have two vascular bundles. 2. Em. navenationofthedisk floretsofm- M is that of the I"apical Composite floller'I (cf. Carlquist, 1961) and is the "Aster-type" or Koch (191)). me vascular supply of the corolla consists of m. fused lateral butadles which extend upward through the limb and divide at the sinuses of the corolla lobes. Afterdividing, theyrunalongthenarginsofthelobesandconverge at t1! spices. ‘11! five stamen bundles branch off the corolla bundles intheregienofthevasculerplensbelowtheapexoftheachene. cm thatch, thests-enbmdlesare MttMnnerthanthecorolla bundles, but,liloetbelatter, theyconsietofa sdnglefileof trachary elnents. The style contains two vascular bmdles which diverge at the Mutation of the ems branches, each bundle supplying astylarla-anchandteuinatingdustbelowtheapexoftbestylar appendages. nintherayaclmes,aaimlebunfleenterstheachnefru thereceptacle. mehmdneerteheenedeteetenuheretreeeer -h8- Plate 8. Fig. l - 6. Details of the vasculation of disk achenes of species of Astereae, viewed laterally. Corolla bundles, 95 style bundles, g; anther bundles unlabeled. Fig. l. Bracmcaae tracmcega, X 80 (299.15; 1.26).. Fig. 2. Dichaetoghora camgstris, X 80 (Prggls M' Fig. 3. Aghanosteghus ramosissimus, X 80 (2e; m 8; m 2&6). Fig. 1;. Townsendia mexicana, I 80 MM). Fig. 5. I. formosa, X 1.0 (m 2219. Fig. 6. g. stLigosa, x ho (was. Armstrog _3_2_0_§_). -50- tracheary elements. in A. m the bundle divides to give rise to four bundles, two lateral, and two dorsi-ventrsl (Plate 7,!‘ig. 2). These may'or>may'not divide in.turn.midway between.base and apext Corolla bundles and stylar bundles branch off the apical plexus. In all other taxa of gtranthium, the disk achenes are two- bundled. 123 single bundle entering Iran the receptacle branches to fans two leterel bundles which traverse the length of the achene. As inthe rayachenes, the twonervesmaybe seenassmallareasof sclerified cells, each directly opposite and external to a bundle. Receptions to the two-bundled condition were noted in sane disk achenes of g. orthflua in which two or three additional bundles were obser- ved (Plate 7, Fig. 5) and in A. eggurucens, in which several achenes had three bundles (Plate 7, Fig. 6). The extra bundles either Join the vascular plexus or branch independently into corolla and anther bundles. In general, t1: achenial bundles rennin free from one another at the plexus, but they are occasionally interconnected. Observations on cleared disk florets of species in other genera of the Astereae were also made. These species are listed in Table 1;. As insggtgggggigg, the majority of species were also found.to have two achenial bundle s, but there were several notable exceptions . It is interesting to note that the species ofmnsted in the Table which have more than two vascular bmdles, also have many- nerved achenes. For maple, g. 392., which Gronquist (19h?) indi- cated as having 8 - lh-nerved achenes, was found to have 12 vascular bundles in several disk achenes. Similarly, g. mtatus has 6-, 7-, or 8-nerved achenes, and I observed 6 bundles in the few disk achenes studied. (I: tip other hand, _E_. ms, which Gronquist Table I... Number of vascular bundles in disk achenes of species of Asteroae. species number of vascular source of material bundles (MSG) Achaetoggron _F_qr__re_ri 2 93 Long & m 1292 A. griseus 2 2131223 gunfire m A. guatemalensis 2 De; £9.13 _6__6§_ _A_. no canus 2 as; £925 631 A. polycephalus 2 Palmr 2 (MO) ghanostephus ramosissimus 1; De; £193 93 lemma 21_6_ seine. 2......"m15 2 M2261 g. szlve stris 2 Botanical Garden, Colmbra Portugal (use 166628) Boltonia asteroides 2 McDonald 2632 Brachycome tmcflcagpa 2 £2933. 126 Chaetopappa asteroides 2 22231-2 gag Dichaetophora campgstrie 2 Linglg £822 m aggentatus 6 Eggs E22 ,3. canus l2 £13}; £2; E. parishii 14 22.42.11 2222 E. vernus 2 133g- 29_0_2- lbwnsendia fomosa 2 m 223 g. moxicena 2 m 1.951 1- 2m 2 2928.9 5229. 2. stflgsa 2 £32392 g ”mating 225 -52- considered to have h-nerved achenes, was found to have two achenial bundles. Although the evidence is not conclusive, it is possible that the number of achenial bundles is correlated with the nuaber of nerves in acne Qecies. It could not be ascertained, however, whether the achenial bundles thaselves or small ribs of sclerified cells consti- tuted the actual nerves of these scheme. Another exception was gmmhus mosissimus in which the disk achenes were found to be h-nerved. Since not all bundles are of equal thickness , it is possible that the achenes of ghanosggphus are cocmonly two-bundled, but that adiitimal bundles are occasionally found. This has been observed to be the case of several mecies of Aetranthim, as indicated above, and it has also been observed in Boltonia asteroides, in which several disk acbnes were three-bundled. B. mstolog l. m £32114. As seen in trans-section, the ground tissue of the laina of the ray florets of Astranthiu consists of loosely arranged parenclwma cells. me vascular bundles are evenly spaced in this ground tissue and lie close to the abaxial epidemis. 'lbe thick- nessofthelninaisuniformthroughout. lbs most conspicuous feature of tie anatomy of the ray lamina is the adaxiel epidermis which consists of a single layer of large cells the outer wall of which is strongly convex and supplied with a thick, shooth cuticle. The abaxial epidermis, however, is made up of a layer of seller cells with end walls which are reunded. lbs thick cuticle is sculptured into pruinent, parallel, sharp-edged ridges and, there- fore, appears Qincse in cross-section. n3 strongly convex cells of -53- the adaxial epidemis are found throughout the lamina. lh the region of the tube, however, the end walls loose their convexity and the cells decrease in size so that the cells of both epidermal layers are alike in appearance. The ray florets of other species of Astereae examined showed no differences in the type of tissues and the position of the vascular bmdles when camared Idlih Astranthiun and with one another. All have the convex adaxlal epidermal cells and the sculpturing of the cuticle of the abaxiel epidermis is also similar. With respect to the thick- ness of the vascular bundles, the lamina of Townsendia stLiggsa, 2. 22m, and '_r_. moxicana have vascular bundles which are considerably thicker than those of any of the other species examined. 2. Q3; Corolla. The five lobes of the disk florets of m- 3.11.593; as in the majority of Comositse, have thickened spices. lb trans-section, these spices are made up of large, rounded cells. Mid- way between the spices and the sinuses, tie corolla lobes are four to six cells thick, including the epidermal cells. From the sinuses dom to the region of the filaments, the corolla wall is only two cells thick, but more call lqers are present around the vascular bundles. In the region of the filaments, the corolla is still two cells thick, but the cells are much larger. Below the level where the filaments diverge fraa t1: corolla tube, the wall is between six and ten cells in tlticlmess. ‘ 'lhe presence of canals associated with the corolla veins is canonly observed. Although sometimes lacking, one, or more cannonly two, canals may be seen lateral to the veins. In A. mthocunoides, -51, only a single canal, which is external to tie vein, may be present. lh all species, these canals appear to run only for short distances, as observed in longitudinal sections of disk florets of A. integ- 29.4.9; subsp. ciliatm. This condition, plus the absence of stained cell deposits, the absence of epithelial cells, and the somewhat irregular shape of tin openings, suggest that tmse canals are air lactmae rather than secretory canals. This also holds true for the other Astereae examined, except for ghancstephus ramosissimus. The disk florets of that species have a single canal located externally to the veins. The presence of epithelial cells indicates that these may be secretory canals. In view of other similarities of m- stgphus to armatic Anthemideae, the presence of secretory canals would not be surprising. 3. galleries. The plyllaries of stranthiisn are scarious- margined and are flattened throughout except near the base where a slight heel is present. The ground tissue of the green midportion consists of spong mesoplnrll which is commonly sclerified. Palisade mesoplvll and secretory canals are absent. Just below the apex where veins are absent, a single pole of sclerenclmaa m be located in the 2 center. lbs ground tissue becomes increasingly sclerified toward the base and the sale is interrupted by vascular bundles. Near the base, where only the midrib is present, the zone of sclerenctvma extends as a continuous band, from one to five cells in thickness, fran near tin midrib to as far as the innermost cells of the scarious margins. The latter are mostly two cells thick, with a single file of one to several cells constituting the very edge. The abaxial epidemic is provided -55- with stomata and trichomes which are lacldng on the adaxial surface. The phyllaries of mhanostgphus ramosissimus, Brennan M- 3325. mohaetophora campgstris, and Iomsendia Pa are similar to those of Astranthium in internal morpholoy. However, the ptyllaries of Bellis perennis, Achaetogeron fleece, lotusendia Mama, and 1. stflggsa are strikingly different. Bellis arennis has ptwllaries in two series which are herbaceous throughout, flattened, and of uniform thickness for the greater portion of their width. 'me ground tissue is made up of sporty mesoplwll which is never sclerified. Palisade cells and secretory canals are lacldng. The ptwllaries of Achastm m are in three to four series ami are heavily sclerified. The outer ptyllaries are relatively thick, with scattered areas of sclerenctyma, whereas the inner ptyllaries are thinner and narrower and are made up of a heavy zone of sclerenclnrma bounded by the two epidermal layers. In the region external to the midrib, some spongy mescplnrll is present. The organisation of the plyllaries of Tomsendia mesicana and I. m is more caplex. The plyllaries are not flattened, but are strongly heeled, the thickened mimortion tapering abrlptly toward the margins as seen in cross-section. me heeled portion is of con- siderable interest in that palisade mescptwll is present on the abaxial side, spongy mesoptyll being present adaldally. At a level corres- ponding to that of the anthers, a continuous some of sclerenclwma ex- tends frm the outer cells of the keeled portion to as far as the prominent scarious margins which are one to two cells thick. lh the narrowed portion of the phyllaries, away from the midrib, sclerified tissue is bordered onlylby the two epidermal layers. m g. m -56- at a level corresponding to that of the achenes, palisade mesophyll is lacking and a continuous band of sclerenctyma traverses the entire width of the ptwllaries, as far as the scarious margins. lb 2. mexicana, at a corresponding level, this band of sclerenclmna is interrupted in the region of the midrib. MORPHOLOGY HABIT. The habit of the species of Letranthimn is closely correlated with their life history. At the beginning of the flowering season the plants are small and commonly have a single, monocephalous stem. All but one species in the genus possess a basal rosette at that time. The exception is A. M, which, at the basis of observations on herbarium material and greenhouse-grown seedlings, appears to lack a basal rosette. As the season progresses, other stems develop, branching occurs, and the plants gain in height. The capitula that are still in bud are typically cernuous, the peduncles elongating and straightening Just prior to anthesis. In most species the basal leaves and lower cauline leaves wither, but they are mostly persistent in _A_. splendens and A. Beemanii. Near the end of the growing season secondary flowering often occurs in the annual species, resulting in the formation of numerous, smaller heads with fewer ray and disk florets. At the end of the season the plants set seed and the camel species die. The perennial species form a new basal rosette and the stems die back and often persist as dried stalks well into the next flattering season. Three species of Astranthiue are annual, one is biennial or short-lived perennial, and six species are perennial. Greenhouse cultures have indicated that the perennial species are capable of coeim into flower as quickly as the amual species. The interpretation of A. menthocomcides as a perennid species -57- -53- requires an explanation. In greenhouse cultures, plants of _A_. xanthoconoides grew vegetatively during the first half of the stunner of 1962 after seeds germinated early that spring. During this tine, single parent plants would give rise to numerous stolons which established new plantlets at their tips. In tin beginning of September the parent plants formed a single flowering aten.after the plants had lost their basal rosette. Following flowering, the parent plants died about the middle of October and those portions of the stolons closest to the parent plants also died. Even though the species showed a pattern of growth typical of other annual species in the genus, it is clearly perennial through stolons . STEHS. The stems of the annual species of Astranthiun are usually nunerous am much-branched above as well as below the middle. The stems of the perennial species are branched to a lesser degree and in a nore open fashion than the annual species. In nost species, the stems are spreading, ascending, or erect, but the stems of the annual A. condinentun and _A_. robustu are often strongly decubent. n:- perennial 5. lastificul approaches being scapose while the perennial A. nnthocolaoides has few to nI-erous stolons in addition to the flowering ate-s. Ste- structure say'therefore provide useful characters for separating species or species groups. Stem color is uniform throughout the genus, varying fro- greenish through greenish- brown to reddish-purple. an stens of all species are grooved and striate, the ribs being of a lighter color than the grooves. Peduncles are conspicuous in all species of Astranthius and are naked except.fbr the occasional presence of one, rarely two, ninete, linear bractlets. where they Join.the capitulua, the peduncles are -59- enlarged, often hollow, and sparsely to densely strigose. Before anthesis, tin peduncles are inconspicuous, the closed heads termin- ating leafy stems. ROOTS. All but one of the perennial species of Astranthium have a fibrous-rootedbbranched or unbranchad caudex that may be vertical, oblique, or horizontal. The exception, A. m has a stout, woody taproot which develops a thick bark. The roots are useful in separating the annual species which are characterised by fibrous roots or slender, light-colored taproots with or without fibrous secondary roots. The taproot of the biennial or short-lived perennial A. 23212- m possesses a swollen stem-root Junction that may be solid or hollow. I LEAVES. The leaves of Astranthium are alternate and are very similar in shape, texture, and pubescence. The basal leaves are the largest and widest in each species and the width is correlated with the width of tin cauline leaves. Thus a narrow-leaved species such as A. orthoMum has correspondingly narrow basal leaves. Since the basal leaves and lower cauline leaves taper gradually toward the leaf- stem junction, there is no differentiation between lamina and petiole. The middle and upper cauline leaves are sessile. The leaf marans of all species are entire, with the exception of A. laetificum which some- times has a few leaves which may have from one to five minute teeth above the middle. leaf gradation, from spathulate or oblanoeolate basal leaves with blunt or nucronate spices to small, linear-lanceolate upper leaves with acuinate spices is striking in the genus. In _A_. M and 5. lactificu, leaf gradation is less strongly expressed. leaf venetion has been found distinctive in A. gloagdm as the leaves -50- of that species are consistently three-nerved in contrast to the one- nerved leaves of the other species. PUBESHCE. The stems and leaves of Astranthium are character- ized by a strigose pubescence. The trichomss are three- to eight- celled and have an enlarged basal cell and an elongated tenninal cell constituting the sharp-pointed apex. The trichemes of the leaves are always somewhat appressed, especially in herbarium specimens, whereas those of the stems are appressed or spreading. The coarsest triohomes are found in the Hinton, Palmer, and Pringle collections of s: m- 299 hrs they may reach 2.0 an in length. INVOLIBRE AND BEETACLE. The phyllaries of Astranthium are flattened and usually imbricated.in two series. The margins are scarious and ciliata or lacerate-ciliate mostly above the middle. The green midportions of the plwllaries are glabrous to densely strigose on the outside and shiny and glabrous within. Phyllary shape ranges from broadly ovate to linear-lanceolate and the spices vary from acute to ciliata-acuminate, often within the same species. Pkwllary length is occasionally'a useful character, battlest species cannot be dif- ferentiated on the basis of phyllary length alone. The receptacle of Astranthiua is'conica'l. It is low-conical in _A_. m but in all other species tends to become steeply conical at maturity. Since Astranthium lacks a pappus for seed dispersal, it is possible that a steep, conical receptacle promotes short-distance wind dispersal by elevating and exposing the mature achenes above the level of the ptwllaries . The surface of the receptacle is alveolate in all species, with a ring of small setae surrounding the achenial attack-lent points. In -61.. mature plants, the receptacle is hollow, the cavity being an upward extension of that of the swollen apex of the peduncle. FLOWS. The ray florets are pistillate and fertile and occur in a single series. Ray color has proven to be a useful taxonomic character when used in conjunction with other characters. The majority of species of Astranthim has white rays, but four species have cyanic rays that are light lavender above and dark lavender below. One species, 5. Bamanii, has rays which are lavender throughout or which are lav- ender above and with alternating bands of light and dark lavender below. These rays nearly always become blue when dried. T13 length and width of the ray corolla is too uniform to be of much value in classification. The tube of the ray is always pubescent with short, blunt trichomes and the apex is usually three-toothed. The style branches are typical of those of other Astereae; one style branch is sometimes considerably shorter than the other but this character is not constant in a species. The disk florets are yellow, five-lobed, and "goblet-shaped”, being abruptly contracted in the threat. The limbs of the disk florets of A. glopgdum are somewhat less contracted below. The length of the disk florets is only useful in distinguistnng A. m from the remaining species of the genus, although thre is some overlap. Author characters are too intangible to be of much value. in apical appendage is always present and the anther bases are rounded. in interestim feature which col-lands attention concerns the color of the erectile tissue of the filaments as observed in greenhouse cultures. With the exceptions of two collections of A. murascens (p; £3 £2}. and 3121), the erectile tissue contains a bright orange pipent before -52- anthesis. This feature may be observed in dried specimens. After anthesis takes place, the erectile sons elongates and the color changes from bright orange to pale yellow. Pollen characters are of little value in the delimitation of the majority of species of Astereae (Wbdehouse, 1959) and this has also been founi to be the case in Astranthium. masurements of the diameter of the grain, spine length, and exine width were observed to overlap to such an extent that separation of the species on the basis of these features was not possible. As in other Astereae,'the pollen of istranthium is spheroidal, tricolpate, and uniformly spinose. ACHENES AND PAPPUS. Achenial characters have been very useful in separating groups of species. In Astranthium, achene shape varies from obovate to oblanoeolate and the achenes are laterally compressed. The achenes of A. glopgdum and A. lactificum are set apart from those of the other members in the genus by their large size, lack of pubes- cence, and the presence of prominent ribs. 5. glopodum has achenes which are two- to four-ribbed and the ribs are lighter in color than the achenial faces which may have up to two conspicuous, longitudinal nerves. The achenes of A: lactificun are two- or three-ribbed but the ribs are less conspicuous than the achenial ribs of A. lo um. The three-ribbed achenes are triquctrous in both species. The remaining members of the genes have small to»medium-sised achenes which are prominently or obscurely two-nerved, lacking true ribs. Sometimes these scheme are slightly triquctrous due to pressure exerted.by neighboring achenes on the receptacle. Pubescent achenes typically have achenial hairs which are duplex, with two connate cells which are rech distally. This type of glochidiate pubescence is -53.. also found in Dichaetophora and Townsendia and underscores the re- lationships of these two genera with Astranthium. The pappus of Astranthium is on the whole inconspicuous and may be present or lacking in the same species or even in the same capi- tulum. Except in A. glopo_dum where it is lacking, the pappus con- sists of a minute, yellowish, whitish, or sometimes brownish ring or crown of setae less than 0.1 mm long. The pappus is most conspicuous in _A_. lsetificum and A. orthopodmn but this may vary even within the same head. Pappus characters are therefore of no value in distin— guishing taxa . DISTRIBUTION AND atoms! Astranthiun is primarily a Mexican genus with eight of the ten species in the genus represented in the montane regions of that country. Tito species are found in the United States, one of which, A' inteegfoliun, ranges southward into the state of Nuevo been in northeastern hexlco. The distribution pattern of the Mexican species is typically 'f-shaped, extending along the Sierra Madre Occidental, the Sierra Madre Oriental, across the Trans-Volcanic Belt of South- Oentral Mexico and southward into the mountains of Oaxaca and Chiapas. No species of Astrsnthiun are known from Central America. The majority of the lbxioan species of Astranthiua occurs between 1700 and 2900 meters elevation, but A. gloadm is feud between 800 and 1500 meters in western lbxico. In addition, 5. Beamanii occurs at altitudes frm 2700 to 3700 meters on Cerro Potosi in Nuevo Iaon where it reaches alpine elevations . Three species are found in western lhxico. The northermost of these, _A_. orthom, occurs in the Sierra Hadre Occidental in Chihuahua and mango, reaching its southsm limit in central mrango. Another western species, _A_. condinentua, ranges from east-central Hichoacan westward an! then northvard into Jalisco. The me of A. E19222! partly overlaps that of 5. condimentum, but the two species are separated fra each other altitudinalJy. In eastern mchoacan, only two localities are known for A. laetificua; one of thee is at ca. 2500 meters elevation in the Transvolcanic alt. Among the eastern species, A. pmurascens is the most widely -5u- -55- distributed. Its range is made up of three populations that are sepa- rated by considerable geographic gaps . The northermost of these populations is in tin Sierra Madre Oriental in an Luis Potosi and mdalgo while tln other two are in the mountains of Oaxaca and Chiapas respectively. Another eastern species, A. xanthocomoidss, occurs in the Sierra Hadre Mental in Bidalgo and in the easternaost mountains of the Transvolcanio son in Puebla, mo... and Veracruz. Although both A. gamma and _A_. menthooaoides are found in Hidalgo, their ranges do not overlap. The remaining eastern species occur in Nusvo Ieon where A. Bea-anti is endemic to Oerro Potosi, principally at alpine and subalpins elevations. Another endemic, g. splendens, ranges from south of Oerro Potosi souttsvard into Oerro Iinadero and cast into adjacent Tamaulipas. That of the lbflcan species of Astranthim are narrow endemios. Three are confined to one state, two species are found in two states, one species occurs in three states, and two are found in four states. All eight of these species occur in montane habitats, but each is separated from neighboring species by geographical or ecological barriers. It is probable that present distribution patterns reflect past changes in both prwsiograplv and climate. All species except 5. M occupy cool, relatively moist open neadowa and open pine-, pine-fir-, and pine-oak forests mostly onbrownsanchlocsandreddishclqlo-s. The habitatofg. m is considerably drier tun that of the otter species, ranging from arid oak forests at 800 meters elevation to semi-arid pine- oak forests at about 1500 meters. Of the Moan species, A. W occupies the most diverse habitats apparently as a consequence of its -66.. wide distribution. In addition to open meadows and pine forests, this species is also found in open Quercus-Taxedium stands and open m- guercus-Juniarus associations. All Mexican species of Astranthium are members of the temperate vegetation zone and belong to the boreal forest- (annual rainfall 30 - 7h inches) and pine-oak forest (annual rainfall 18 - 70 inches) conununities (cf. Leopold, 1950). The United States species Astranthiuu integgfoliuu is the geo- graphically most widespread member of the genus; it is here recognized as cmprising two subspecies. Subspecies ciliatum ranges from Nuevo leon, lbxlco, where it reaches elevations of 2500 ft. through east- central Texas northward into Oklalma and the southermost counties of Kansas and Missouri and eastwwd into Arkansas where it overlaps with subsp. integggoliua. In Texas, subsp. ciliatum occupies six different vegetational areas at elevations ranging from sea level to about 800 ft. (Gould, 1962). Although differing in species cmpositicn, tines vegetational areas are predominantly characterized by acid sandy or as” loan soils. On the basis of distributional data obtained from herbarim labels, subsp. ciliattan has its greatest concentration in the Gulf Prairies and Marsha, Post Oak Savannah, and mackland Prairies vegetational areas, with average annual rainfall ranging frat I) to 50 inclns. m Oklahma, _A_. integgoliu subsp. ciliata is confined to the eastern half of the state. a: tin basis of distribution data, subsp. My occupies four major vegetation zones as outlined by Bruner (1931). the true prairie and oak-hickory savannah associations in central and cast-central Oklahoma, and tin eastern grassland and oak- hickory forest associations in the eastern part of the state. -67.. Corresponding average annual precipitation increases eastward, ranging from 30 inches in the central portion of the state to 1.5 inches in the east. The eastern part of the range of A. integifolium is occupied by subsp. integifolim. This subspecies ranges from Arkansas, where it overlaps with subsp. ciliatum, eastward to Tennessee and Kentucky and southward into central Mississippi and the northern parts of Alabama and Georgia. Available collection data from herbarium specimens in- dicate that this subspecies occupies calcareous soils in relatively dry to moist, open to shady woodland habitats. The last species under consideration is A: robustun which is a narrow endemic found only in the Trans-Paces region north of Big Bend National Park in extreme western Texas at elevations between 2900 and 13833 ft. 5° robustun occupies habitats ranging from arid pinehill flats and open, arid roadsides to arid canyons in the mountains on soils that are predominantly alkaline. Ecologically, A. robustun is a member of the "Trans Pecos, Mountains and Basins” vegetational area of Gould (1962), which is characterized by an average annual rainfall of less than 12 inches. Although the region in which A. robustun is found borders on Chihuahua, the species has not been reported fra Mexico; it is also unknown from Big Bend National Park. EVOLUTION A. m. Since gtranttrium is primarily a Headoan, montane genus re- stricted to the Sierra Hadre Occidental, the sierra Madre Oriental, the Transvolcanic Belt, and the nomtains of central Oaxaca and central Ohiapas, it is thought that the evolution of the genus is closely correlated with the plvsiographic history of these mountain ranges. Schuclnrt (1928) suggested that the present relief of Mexico was the result of elevation and block faulting in the late Pliocene and Pleistocene. Theyer (1916) indicated that the Transvelcanic th owed its relief to vulcanism starting in tin Pliocene. White (1962) v considered the oldest volcanic rocks exposed along the lowermost western slopes of Iztaccihuatl to be of mid-Thrtiary age. According to Garfias and Chapin (19159), the Sierra Hadre Occidental was up- lifted in the Miocene, the Sierra Hadre Mental in the Pliocene, and the Sierra de Ohiapas in late Pliocene. nae present high relief of ,much of Mexico is thus considered to be of relatively recent Main. During the tertiary, concomitant with erogenio events, there was a gradual, worldwide trend toward cooling and during (mired, 1960), cullinating in tln great ice advances of the Pleistocene. m (1956) postulated that during the Pleistocene, '...there appears to have occurred at that period of naximn glaciation a clinal depres- sion of mean-temperatures that amounted to 5° 1‘ at the equator, to looratlatitudea 35°tohO°N, andt025° atthe edge oftbice -68- .69. sheet". Decreases in water temperatures as well as in the level of tin oceans were also evident at that time (nun-m. 1955; Flint, 1957). Climatic cooling durirg the Pleistocene ny be inferred from the {resenoe of glacial features on Iataccihuatl at elevations as law as 21:50 n (White, 1962), indicating an sltitudinal difference of about 2000 mters bemn existing and former glaciers. Additional evidence was presented by Lcrsnso (1959) who noted the presence of glacial features on mountains which do not have glaciers at the present tine. Sears and Clisby (1952), in a shady of soil-case nu Mexico 01w, shared a correlation betteen'oontinental glaciatim an! the capaci- tion of fossil-pollen profile s, as indicated by the memos or absence of pollen of upland species of trees at difth depths. may concluded that "....moiet and/or cool conditions represent periods d ice advance, whereas dry aid/(r warm "meant intervals cf retreat”. In Cmtral America, past cooler clintes are indicated by evidence of glaciaticn on Cerro Chirippo in the Sierra Talannca, (Heyl, 1955), and by polls: pofiles suggesting major demssions of vegetation sense in Costa Rica (Martin, 1961). Martin (1958), sunmarising snxdromental changes during the Pleistocene in western North Mica and Central America, postulated a M110 to 160) foot downward displacunt of biotic some in the soufliwest during the period of maximal gladation. On this basis, Bab:- and (rear (1962) suggested that in response to Pleistocme climatic cooling, tenpsrate plate and ani-ls were able to migrate alum the Sierra Hadre Occidental which served as a chain-like avenue 1' diapreal. Darin; the Plastooeu, the Sierra Hadre Oriental, the Sierra Hadre del Sur, and the mountain of neural Oaxaca and central Ohiapas met also have -70- served as principal migration routes for temperate biota. Although.temperate plants may have existed in.Hexico since early Tertiary or even Cretaceous times (McVaugh, 1952), it has been sug- gested by Sharp (1953) and Dressler (19514) that the Pleistocene was the most favorable time for the dispersal of temperate plants. Similarly, Raven (1963) indicated that dispersal of the majority'of temperate disjuncts in North.and.South America may have taken.place during the late Pliocene or Pleistocene. With.respect to Astranthium, Pleistocene dispersal and.evolution of the genus is strongly suggested by several lines of evidence: (1) the genus is of limited distribution and.near1y all species are endemics; (2) species which are related on the basis of morphological characters often occupy adjacent ranges; (3) new species have some genic material in cannon, as indicated by artificial crosses, suggesting recent divergence from ancestral stock. This evidence is especially impressive in the light of the fact that in rapidly evolving herbs such as ‘gtranthigg, chromosomal differ- ences may build up rapidlyt It seems most likely that the ancestral stock of Agtrenthium originated in west-central Mexico where the most primitive species, _A_. ylgodum, is found. The range of tin next most primitive species, A. lastificuu, is only about 150 miles to the east. n10 momhological gap between the primitive qaecies with chrmosane timbers on a base of g - 5 (A. M and A. lastificua) is considerably greater thanthegapbetweenawpairofqeciesinthex-handx-Beeries. Agtggnthdunrgylggodumumay thus be considered to be older than the other'apecies. Partner-ore, it occurs in.relatively dry, lowreleva- tion habitats which.have been.in.existence in.Hhxioo during:amchxof -71- the Tertiary. oh this basis, the Pliocene seems to be a likely time during which A. ylgodum might have originated. As Tertiary orogsny and vulcaninl increased the availability of temperate montane habitats, some populations of A. glopodun may have moved into these scattered and biologically open habitats. The closely related 5. m may have originated directly from.these populations. ‘Hhen additional cool, montane habitats became available during the Pleistocene, stock of Astranthium was able to spread along mon- tane avenues of dispersal across Mexico. This may have taken place during each of the cold.periods of the Pleistocene. Increasingly warm.inter~ and.post-glacial conditions may have been responsible for disjunction and further evolution. It is possible, that the last waning trend gave rise to the present-day species. During the last, or perhaps one of the earlier wane, dry periods, some stock of Astranthiuu migrated northward along the Sierra Madre Oriental into the Trans-Pecos region of Texas and became specialised.as a result of adaptation to iseric conditions, forming A. robustum. Closely related stockaigrated.ncrthnard into the southpcentral and east-central mited States and became differentiated into two subqnscies. (he of A these, _A_. integiiolium subsp. ciliata, appears to occupy somewhat drier habitats than does the other, subspo integgifolinn. The dis- tribution pattern of the genus indicates that geographic isolatial has played an important role in the evolution of the species. This appears also to be the case in the closely related.genus Townsendia. However,'where no sexne1.podyploids are known to exist in Townsendia (mean, 195?), P°1JPloiw is an additional factor which may have played a role in the evolutionary'pattern.of égtrenthium. -72- lists for self-convatibility in Etranthiun have indicated that all species are self-fertile . Self-fertility greatly facilitates migration since a single achene, landing in a favorable locality, is potentially capable of forming a new pqaulation. In W migration was likely accomplished by relatively short-distance Jumps, since the achenes lack pappus barbs or bristles for either animal or find dispersal. It is possible, however, that some wind dispersal takes place, since the conical receptacle elevates the small achenes above the pkvllaries, allowing them to be detached by gusts of wind and blown perhaps for considerable distances. The Isthnus of Tehuantepec, which separates the Oaxaca and Chiapas populations of Astranthiun pguraecens, (cf. Plate lb), may serve as an maple of an area which may have been crossed by m- Eggs; in a single Jump. The presence of tropical Miocene fossils (Rm, 1923) indicates that this area was covered with trqaical vegetation during much of the Tertiary. The dispersal of these trop- ical species was apparently uninterrupted, since evidence for the absence of Cenozoic seams across the Isthnus is now well doomented (Durham _e_t_ 31;, 1955). It also appears that the Isthnue has always . been an area of low relief (aahuehert, 1935) and even today the highest elevation is a mere 825 ft (Baker, 193)) on the continental divide. Although Stuart (1951) mated that dining the Pleistocene, "...a relatively cool enviroment must certainly have fouled a bridge across the relatively low lath-us of iehnantepec', it is doubtful whether climatic conditiaia were ever cool enough for temperate plants to exist in the area. For angle, a domard depression of vegetation sales of 1000 to 1500 ft, as postulated for more northern areas by .73.. Martin (1958), would be insufficient b establish a cone of temperate vegetation on the lemma. It seems highly pobable, therefu'e, that the lemme of Tehuantepec has always been tropical, em during the tins of maxim glaciation, and that temperate species such as Aetranthitm masonic crossed this tropical gap by a single Jump. Analyses of pollen profiles could be of great interest in this respect, but such studies have not yet been carried out in the Isthms of Ibhuantepec. s. Evolutionag m. ‘ 1b interpetation of evolutionary trade in Aetranthiun is based on three principal kinds of indirect evidence. (1) here are morphological (and libly also physiological) specializatiom which may be inferred frat dish'ibution pattern and ha‘oltat requireuents of the species. (2) Sililar changes have taken place in several characters in both ietranthiun and related genera, and these appar- ently are correlated. (3) 'Ihere has been a general tendmcy for reducflon in various pub of the plant body. Primitive and advanced curactere in Ashanthiun (cf. Inble S) inferred has! the disbibutim and habitat. involve print-11y the per- emial habit an! the woody taproot or fibrous roots. me species thich have these diaracters occupy relatively neeic areas within the range of the genus, whereas the annuals with slender taproots occur in drier habitats. ibis trend my also be inferred fro the phydographic history of the areas in which Astrmihinn occurs (see undc Origin). Sililar Table 5. molutionary trends in Astranthiua. primitive characters advanced characters l. 2. 3. perennial habit woody taproot little gradation in length and width of cauline leaves few, large bade long phsllariss plwllaries with narrow scarious margins large floral parts large achenes glabrous achenes achene with 2 or 3 con- spicuous ribs achenes with more than two vascular bundles basic chm-cease nmber ; - 5 diploid? large chromosomes symetrical karyotypes annual habit fibrous-rooted caudices to slender taproots strong, upward gradation in length and width of cauline leaves nunerous , relatively small heads short plvllaries plyllaries with wide scarious margins auller floral parts small. achenes glochidiate-pubescent achenes achenes 2-nerved achenes with 2 vascular bundles basic chroncsue number 5 - 3 9013910165! small chronosues asymetrical karyotypes -75- interpretations have been applied to new other groups of plants. A comiderable nunber of evolutionary trends in gtranthium are apparently correlated. Correlated resemblances are indicative of plvlogenetic relationships (Ownbey and lace, 1955), and by the same token related species subjected to similar enviromental stresses are likely to undergo parallel evolutionary changes. The species Idth primitive root structure also shows little gradation in leaf size, and it has few, large heads, long plwllaries with narrow scarious margins, large floral parts, large, glabrous achenes with two or three conspicuous ribs, the highest basic chromosome nunber, the largest chromosomes, and the most syrmetrical karyotype. In addition, the most primitive species of Astranthium has more achenial bundles than arw other species and this is considered to be a primitive character (of. Carlquist, 1961). Bsbcock's (19142) studies on _Cr_8Lls_ provides additional support for the interpretation of the higher basic chromosane nunbers, the larger chromosome, and the symmetrical karyotypes as primitive characters. The most primitive species of Townsendia also has almost all of the-same characters (of. Beaman, 1957) as those which are interpreted as primitive in Astranthium. The trends of specialization in Townsendia are also similar to those in istranthinn. The species of ' gtranthiun which some to qualify as the most primitive species of the genus is i- glgpodun, it is characterized by all of the primitive characters listed in Table 5. It does not necessarily follow, however, that because certain char- acters occur in a species which is primitive, that all characters are primitive. In a genus as relatively young as Astranthimn (see under Origin), however, there is a greater probability that most -7e- primitive characters have been retained than may be trm in relatively older genera . The trends in specialization almost all involve reduction. the slander taproots of the annuals is presumably reduced from t1! woody taproot qr fibrous roots of the perennials. (radation in the cauline leaves is the result of reduction of tin Imper and middle leaves. Tb change from large heads to small heads, long to short plwllaries, large to small disk florets, large to small achenes, ribbed achenes with more than two vascular bundles to nerved achenes with only two vascular bundles, fromg-Stcg- 3, andfromlarge to emallohromo- somes are all features of reduction. Very few features of elaboration are evident in ggmthiun. Pclyploidy is perhaps the most obvious of these, and the change fru few heads to nunerous heads, and from glabrous achenes to glochidiate- pubescent achenee may involve elaboration. me latter trend is also evident in Townsendia. Evolutionary change through reduction probably has a greater probability of occurrence than change through elabor- ation (Stebbins, 1950). It is likely that in a recent and rapidly evolving genus such as Astranthiu features of reduction would be more frequent thm those of elaboration. ~ TAIDNOHJB CONCEPTS In Astranthium, I have recognized as species tane which are .morphologically distinct at all times and.which are isolated from one another by various barriers to gene exchange. These barriers are primarily geographical and ecological. All species in the genus are allopatric and no interspecific hybrids are known. This would sug- gest that the present-day species of gtranthiua hove achieved their distinctness while being spatially isolated from.each other. when species are crossed experimentally, partial or complete sterility barriers become evident . Homoploid hybrids have reduced fertility possibly because of minute genie or structural differences in the chromosoles. On the other hand, heteroploid hybrida:nay'be completely sterile, especially when.parenta with. widely different chromosome numbers are crossed. these partial or complete genetic barriers are present both at the diploid.and. polyploid.level. Where chronosomal races differ from one another in a few, but constant characters, and, in addition, are separated‘by geographical and ecological barriers, 1 have considered these to be distinct species. The hexaploid A. gglendens and the octoploid A. Beamanii may serve as examples in this remect. I have recognised as subspecies geographic races which differ morphologically'in.relatively constant, mostly quantitative characters, uhich.occupy considerable ranges, and.whioh.intergrade in the area where their’renges come into contact. -77- GENERIC REIATIONSHD’S The generic relatives of Astranthiua are not revealed by the nomenclatural history of the species in the genus. Host taxa treated by Larsen (1933) and accepted in the present nonograph were trans- ferred by her from worse” A. xanthcconoddes and A. m which were in 3mm” and Loggia. respectively. Although £113 and Astranthium were long considered as con- generic, the two genera have little in colleen. The species of £113 are scapose and have few, relatively wide, herbaceous, biseriate ptwllariea with blunt spices, and large, mostly glabrous, ribbed achenes . Astranthiun species are caulescent and have nmaeroue , scarious-marched, noetly bdseriate plvllaries with acute or acuminate spices. Mthemre, the distant geographic separation argues against any close ties between tin two genera since §_l_._li_s- is primarily found in the Mediterranean region, although adventive in North America (a. m~ Arnott (in seem, 1835) mas-steel that 9323 nemesis (istranthlu integgcnuu) was the as. species as been“ mtbcco-oddes (Astrenthium xenthocaoi ). It is perhaps for this reesm that Bcffnenn (1891;) placed istrenthiun in synonym under Ragga-Io. In her monograph of the Australasian Reggae, Davis (19%) concluded on the basis of a study of achenes and phyllariee that w Astranthiue and nuance» originated from a cannon ancestcr, with iatrenthium and Broom” the closest of the three -73- -79- genera. Although Bracflcome resembles some species of Astranthium in the shape of the achenes, the most primitive, scapose, fibrous-rooted and small-headed species of that genus are very different frau the caulescent, taprooted and large-headed 5. m, the most primi- tive species of Aetranthim. Furthemore, the leaves of Bremen“ are cosmmly toothed or variously dissected whereas the leaves of Astrenthiun are almost always entire. Since distant geographic separation is an additional factor that argues against any relationship of Astrenthim with either §e_l_li_a_ or Bracmome, it seems reasonable to suggest that the generic relatives of the genus should be sought among North American or Helicon Astereae. Larsen (1933) thought the genus to be related to _I_e_e_r_l_i;s_, Achaetgggron, and iphanostephus. The genus [eerlia is not in Ghaetppappe (Shimers, l9h6a) which is well set apart fran .Astranthim by the often high and narrow involucre, the unequal galleries in 2 - 6 series, the flat or slightly convex receptacle, and few- to many-nerved, compressed or subterete achenes. alimere (l9h6b) outsiders Ghaetmma to be re- lated to Leucelene. The relationships bottleen Astrenthipm and Achaetggeron are not apparent follating the removal of Astranthiun medcenin and 5. gusto-slenee. m the other hand, Acheet_o_ggron is most closely related to m and may actually belcng in that genus (De Jong and Mrs, 1963). A relatimship between Aetrenthium and Westephus is indicated by the conical receptacle, the scarims-nergi.ned planaries, and the basic chromosome nuebers of g- 3, h, and S which the two general have in comm. monasteaa is unlike Aetrenthiua, however, in the unequal, charteceous pulleries in 3 - 1: series, and curved, columnar achenes as well as leaves that are toothed to deeply pinnatifid in -30- most species. Sinners (l9h6c) included Astranthim in an arrangement of genera of Astereae a: the basis of the conical receptacle that these genera have in common. Of these genera, Astranthiua, nichsetghora, and Boltonia fall into one group on the basis of key characters. Beeman (1957) cmsidered Townsendia also to be a member of this group because of the conical receptacle in 2. fomosa. Be inucated that Townsendia and Dichaetgmra were the closest members of this alliance, with Aetrenthium saewhet more distant and Boltonia further removed. A stuw which I have node of the four genera suggests that istrenthiun and Townsend: are even closer than previously indicated by baman. A close relationship between Astrenthiue and Townsendie is indicated by the nature of the achenes which are obovate or oblanoeolate and often glochidiate-pubescent, by the scarious-margin“ plyllaries and the conical receptacle. he leaves of both genera are entire, spsthulate to linear and strmgly reduced upward and have siniler pubescence. Furthermore, Astrenttmn orthom has a swollen stu- rcot Junctim which is matched in several species of Townsendisand lavender rays are found in species of both genera. Most important is the fact that both genera are naitene and their centers of diversity are not far apart in the cordillere of the southern United States and Race respectively (of. Beanan, 1957). This would suggest that the two genera have had a somewhat einiler evolutimery history, although apdns, canon in Townsendia, is unknown in Aetrenthiu. As in Townsendia, latitudinal and eltitudinel influences are iaportant in the distributiai of the mmtene species of Astranthim and narrow caddie- prevails in both genera. m the whole, Astrentlliua requires -31- more nesic conditions than Townsendia, although 5. rchustun parallels the more specialised species of Townsenda in its arid habitat and alkaline soil requirements. Hcrphclcgically, Townsendia appears to be less specialised than Astranthiun in the relatively wide, 2 - 7- seriste phyllaries, the large number of ray florets in sale species, and ribbed achenes. Astranthim is also related to mohaetghora (or. gunners, l9h6c) . In addition to the cortical receptacle, the two genera have in canon equal, scariousa-nargined plwllaries in two series, disk florets which have the limb abruptly contracted at the throat, and glochidiate-pubescent, cupressed achenes. In habit, nichaetcphora canpestrls is very close to A. intgg_li_f_cliun, especially subsp. ciliatun. nichaetflora differs from Astranthiun, however, in the presence or! a snall but conspicuous squanellate pappus and promi- nently Hinged achenes winch indicate a relaticnship With Boltonia with which the genus was tonerly united by Bentham and Becker (1883). Boltmia is clearly related to £393. and m and is distant Iran Astranthiun. Further references to the generic relatives of “tram-him nay be found in tin section at the subtrlbal positim of the genus. TI-E SUBTRIBAL POSITION 01" “mm: THE DISPUSI‘I'lON OF THE am 01‘ I'HE ASTERELE-BMINAE Astranthiun has been assigned conventionally to the subtribs Bellidinao (in synonym \mdsr Bracfigaae) by I-bffiaann (1891i) and was considered to belong to tin "Dollie-type” by Bentham (1873). 'me subtribs is separated tron the closely related subtribs Asterinas mainly on the basis of the reduced pappus. Hermann considered this separation to be artificial and Shinners (191:9) indicated that the subtribs Bellidinae itself "... as defined by Hermann... is decidedly heterogeneous and artificial”. Since Astranthiun has been found in this stuw to be more closely related to genera in tin Asterinae, a survey of herbariun material of the remaining genera of the Bellidinae was undertaken in order to assess their relationships. '11: results or this survey fully agree with Shinners' observations and indicate that these genera are not at all, or only distantly related to each otlnr and that they have closer congeners elsewhere on the basis of characters other than those of tin pappus. With reference to the generic relatives or Astranthim, it has been pointed out that the genus has no close relatives in the Bellidinas but is closely related to Wendie and mohaetoghora in the Asterinae. this close relationship certainly supports the place-tent of Astranthiue next to these genera in the uterinae. 01‘ the New fibrld genera of tin Ellidinae, Keerlia is now in -82- -83- Ghaetcpama (gunners, l9h6a) and m has been placed in the sub- tribs Grangeinae near arms (Shinners, 1910). I concur fully in the disposition of these two genera. The genus Greenella was considered by Selbrig (1960) to form a close alliance with W mummy, Guttierreaia, and Ethocemlm in the Solidagininee and is well-placed in that subtribs, differing mainly in the color of the rays. The close relationships of the thrican genus AchaeMron with m were discussed by Cronquist (19h?) and more recently by De Jong and nengpre (1963) end the genus is certainly to be placed next to m in tin Asterinae. The genus Qhanosteghus, related to Astranthiun and other genera of the Asterinae having a conical receptacle, is not out of place in that subtribs if grouped with these genera. Of the remaining genera, the European gill}; has no close rela- tives in tin Bellidinae but is most closely related to £113; and Bellidiastrua in the Asterinae with which it m be placed on the basis of the scapose habit, spathulate, toothed or entire radical leaves, herbaceous, biseriate pivllaries, and ribbed achenes which the three genera have in canon. '33 Australasian genus EacMone often has pappus bristles which are no more reduced than those of new nenbers of the Asterinae and in all other characters the genus is clearly a leather of that subtribs. It has no close relatives in tin Bellidinae, whereas it is related to 933313 and ninuria in the Asterinae (Buthan, 1873). It is suggested, therefore, that Bracgcaee be placed with these two genera. m. Owenda Davis, who has nonographed both m and Gelotis, concurs fully in w disposition of m (personal coununicaticn) . -8h- The genera Menommum, Moth, Laggnoghora, and Solenom are related on the basis or achenial characters, plvllaries which are mostly chartaceous and scarious-nargined, and ray florets which are often greatly reduced. The scheme or the tour genera are similar in shape, texture, and nervation and often have a conspicuous beak which may be produced into a sticky, glandular ring. In some species or griactis and Laggnoghora this beak is reduced and the glandular ring is essentially sessile. mmsamm and Meetis are caulescent and have ray florets which are mostly nultiseriate and nuch reduced, often tubular. 'rhe ray florets of genome” are in l to few series and are also reduced although never tubular. me genus salmon! is considered to be derived from W by Davis (1950), differing mainly in the tubular or very short-upped ray florets. The canbined characters of the four genera are typical of. the subtribs Orangeinae and it is suggested that these genera be placed in that subtribs. This transfer also receives support from the fact that Bentham (1873) re- ferred to museum and mucus as belonging to the "Oranges- type". The generic relationships of the remaining genus, Erodiomllm, are less obvious. The genus is unique in its nultiseriate pistillate florets, the outer of which are radiate and which may have 2 or 3 tree stamens. The corollas or the inner pistillate florets are reduced to a densely pubescent tube less than 1.0 an in length and are subtended by stifi', men, receptacular bracts with sharp points which become reflexed at maturity. niece bracts are not present anong the disk florets. Although the style branches or the sterile disk florets lack stigaatic surfaces and are used for the greater part or their length, Table 6. Proposed disposition of the genera of the tribe Astereae, subtribs Bellidinah. genera of the new subtribal position in Ballidinae the tribe Astereae Ercdioggllum Asterinae i—L‘ W" h“ 322:3: ;.?°::.——a”2:i‘::$"m¥‘ Greenella Sondauninae, with An hi s flhiacmh Guttierrezfi @- as... m and m Laggnoghora Grangeinae W Gram-inns MMsamum Grangeinae ctis Grangeinae Achaetoggron Asterinae, near m Brae cone Astezdnae, near 993119. ani Iflnuria 2%- exmwww Astranthiua Asterinae, near W and MohaeMa -35- they are typically Asteroid, being glabrous on the inside and pubescent on the outside. 111s well-developed cyanic rays of Mogflllm would suggest that the genus is a nunber or the subtribs Asterinae, notwithstanding the multiseriate, greatly reduced inner female florets which are reminiscent of the Grangeinae. The determination or the exact plnrlstic position or their genus in the Aeterinae must await further study. MOLUDED SPECIES Larsen (1933) placed in Astranthium species which were similar to the type species of the genus in the reduced or absent pappus, but which differed greatly not only in other floral characters, but also in vegetative characters. The taxa which are anomalous in the genus on this basis are Astranthiun m (Blake) Larsen, A. mezlcanum (A. Gray) Larsen var. mexicamum, A. mencanum (A. Gray) Larsen var. chihuahuense Larsen, and the more recently described A. ggatemalense males. These four tane have the following floral characters in cannon: (l) A flat or slightly convex receptacle; (2) Numerous, un- equal, narrow, herbaceous or subherbaceous ptwllaries; (3) Numerous (more than 60) narrow-rayed pistillate florets in more than one series; (h) Oblong, two-nerved achenes which are often widest about the middle and taper off towards the base and apex. he leaves of A. mezicanun var. mexicanum are entire to coarsely toothed while those. of the other taxa are entire but differ from typical members of Astranthium in shape and texture. The floral characters listed above, together with the reduced or absent pappus, are characteristic of the genus Achaetggrcn. The status of this genus was recently 41......“ by n- m and Inngpre (1963) and it was provisionally considered to be distinct by these authors. Bendix; further studies in Achaetoggron, the above ten are best placed in that genus. me following transfers and name changes will be effective: -33- 1. Achaetogrcn guatenalensis (Blake) De Jong, cash. nov. gtranthim mtenalense Blaine, Brittonia 2: 335. 1937. Ups: GUATEMALA: in cpen pinewoods, along trail between mehuete- nango and Selena, Sierra Cuchnnatanes, apt. mehuetenengo, alt. 3335 m (11,000 ft), 15 Sept 1931:, gate}: £2.22 (helotype. an; isotype, 1n). 2. Acheetoggron mencanus (A. Gray) De Jong, comb. nov. wmncam A. Gray, Smiths. Contr. S: 93. 1852. Bt‘acficome nexicana (A. Gray) miche, Fl. Ear”: 181;. 1926. gtranthium madcanun (A. Gray) Larsen, Ann. no. not. Gard. 20: 32. 1933- Iype: sum or MEXICO: without locality, 26 April 1819, m 1.0}. (holotype, 0m; isotype, 140:). Beth A. guatemalensis (P. - 9) and A. mexicanus (9. r 9, 18) have chromosome numbers on a base of 5 - 9, which is also the basic chromosome number of Achaet_c_>mron (De Jong and Iongpre, 1963). A. mexicanus bears a remarkable resemblance to Achaetoggron Geleottii and A. affinis, but insufficient material of tin latter two did not allow for a critical and detailed comparison of the three species. More detailed studies may show them to be conspecific. 3. Achaetoggron arennis De Jong, none. at stat. nov. Astranthiun nexicanln (A. Gray) Larsen var. chihuahuenee Larsen, Ann. 80. mt. Gard. 20: 32. 1933. ‘lype: CHIHUAHUA: canyons of the Sierra Hadre (Occidental), it Oct 1888, my; (holotype, H01; isotypes, G31, 2 sheets; 1133,13 Pm). -39- This tamer: is sufficiently distinct frat A. mexicanus in its entire leaves and glabrous achenes with a small, but conspicuous pappus crown to warrant its recognition as a species. Since an earlier Aghaetoggron chihuahueneis was described by Larsen (in Blake, 191:0), a new epithet has been coined. h. Achaetgmron 9.11M (Blake) De Jong, comb. nov. Maw Blake, Contr. 0.3. Net. Herb. 22: 591:. 1921:. Aetranthiun m (Blake) Larsen, Ann. no. Bot. Gard. 20: 30. 1933. Type: DURANGO: I) Die N of Guanacevi, alt. 2M0 m to 271:5 m, 18 Au 1898, m m (holotype, 0513 isotypo, om). Achaetogron m_i_1_aA:_s_ is only known from tin type locality in northern Durango and is distinct in its large, obovate-oblong leaves with long, slender petioles, linear-lanceolate cauline leaves which are reduced upward, and few, large, many-flowered heads. SYSM'HG 'IREA'MENT 'lhe herbaria Irma tdzich specims were borrowed are listed belat. Abbreviations are those in thefifth edition (1‘ "Index Herbariomm" (Lardner and Starlet, 1961:). Chicago Natural Hista'y Huseum (F); Cmservatoire st Jerdin Boteniquee, Geneva (G); Gray Herbarium, Harvard University (GI-I); University of Glasgow ((1.); Univerdty of Kansas (KANU); University of Michigan (MICK); Missouri Botanical Garden ()0); Michigan State University (1450); University of Oklaha- (on); Oklahau State Uni- versity (0m), mseum National d'Histoire Naturelle, Paris (P); Philadelphia Acedeuw of Natural Sciences (PH); Southern Methodist Unvereity (SHU); Sul Ross State College (SRSC); University of beas (EX); University of Califcrnia (00); U. 3. National Rhesus: (vs); University of Wisconsin (WIS). AS'IRAN'IHIW Nutt. Trans. Am. Phil. Soc. N. S. 7: 312. 1.81:1. Amual, biennial, an! peremdal, caulesomt herbs with taproots, fibrous roots, tr fibrous-rooted caudioes; stem erect, ascending, or dscumbent, striate and grooved, sparingly to densely strigose with spreading to appreesed frictions; basal leaves broadly spethulate to narrowly oblanoeolate (lacking in A. M), essentially glabrous to somewhat densely appressed-strigoee; caulire leaves alternate, sessile, speflmlate to linear, mostly strongly reduced upward, glabrous to densely strigose, the urgins entire, rarely tootlnd, ciliate, st.- tinis revolnte, the apices moronate to aouminab; heads mdium-sised -9o- -91- to large (secondary heads often mall), cernuous and subtended by the upper cauline leaves prior to anthesis, cos-only long-pedunculate, the peduncles naked or with one or two mall, linear, leaf-like bracts; receptacle conical, rarely low-conical, alveolate; involucre caspanu- late orlhmnispherical; phyllaries inhriosted.in.mostly 2, rerely 3 series (sometimes a single series in secondary heads), flattened, broadly ovate to linear-lanceolate, glabrous to densely strigose on the outside, glabrous and shiny within, the margins pruninently scarious, rarely near-herbaceous throughout, lacerate-ciliate mostly above the middle, the spices acute or lacerate-acminate; ray florets pistillate, fertile, in one series, 10 - 35, white throughout, or white above and dark lavender'below or the lower surface with.alternating bands of dark and light lavender, the upper surface appearing light lavender when viewed frm above, tb tube with scattered hairs, the spices 2 - h-dentate or -denticulate; disk florets perfect, to - 26G, yellow, the corolla rarely'slightly, but more commonly abruptly contracted.at the minutely pubescent throat; anthers with.apical appendages and rounded anther bases, the erectile tissue in the filneents nearly always of a bright orange color prior to anthesie, changing to pale yellow at anthesis; style brenclus of the disk glabrous on the inside, pubescent without, the stylar appendages ovate to ovate-lanceolate, slightlyaherterthantoabmtoneandahelftdnestblengthofthe stigmatic lines; achenes obovate to oblanoeolate, laterally'coepressed, glabrous or glochidiatespubescent with.duplexs bifurcate hairs, greenishs bronm to dark brown; disk and ray achenes 2-nerved or obscurely so, or conspicuously 2- or 3-ribbed, triquctrous when 3-ribbsd, the faces of the ribbed achenes nervelese or with one or two prminent nerves; -92- pappus absent or a minute crown of a few whitish or yellowish teeth less than 0.1 m high. 'lype species and originally the only species in the genus: Astranthim integgoliua (Hichx.) Nutt. A. A. B. B. Key to the Species Achenes 2- or 3—ribbed, triquctrous when 3-ribbed, the achenial faces with l or 2 conspicuous, longitudinal nerves or nerveless, glabrous, 2.5 - 3.2 m long; cauline leaves nearly similar in size and shape throughout; rays white; perennials .................. B Plants with a stout or slender W taproot, often with rough bark; cauline leaves 3-nerved; limb of the disk florets 3.1: - h.1 m long, slightly contracted at the throat; outermost phyllariee 6.0 - 8.5 m long, ovate-lanceolate to linear-lanceo- late; acmnial ribs of a lighter color than the faces; achenes nerved or nerveless; pappus absent 5 plants frat Nuarit, Jelisco, and lflchoacan................................. l. A. M Plants with a fibrous-rooted caudex; cauline leaves sometimes with l - 5 small, callous-tipped teeth; limb of the disk florets 3.3 - 3.7 mm long, abruptly contracted at the throat; outermost phyllaries h.S - 6.0 me long, broadly ovate to ovate-lanceolats; achenial ribs of the same color as the nerveless faces; pappus absent or a small, whitish crown of teeth lees tinn 0.1 mm high; plants from eastern lfl.choaoan................ 2. A. lastificun achenes 2-nerved, never ribbed, glabrous or glochidiate-pubescent, 1.8 - 2.3 II long, epappose or with a clan, whitish or yellowish crew of teeth less than 0.1 use high; cauline leaves strongly re- duced upward in length and width; ms white, or lavender below and pale lavender above; annuals, biennials, and peremials.... 0 -93— -93- 0. Rays white, sometimes roseate in drying, but never bright blue or purple when dry ....................................... D 0. Plants with fibrous-rooted caudices; plants fro: eastern lbxico....................................................... E E. mot, cpen-branched perennials; flowering stone numerous, branched at or below the middle; plants fraa in Luis Potcsi, Eldalgo, Oaxaca, and Chiapas............ 3. A. pmursscens E. Procumbent, stolcniferous perennials, the stolons basal or cauline; erect flowering stems rarely more than 1; plants from mdalgo, Puebla, Tlancala, and Venom"..................... ....................................... h. A. zanthocasoides D. Plants with short or slender taproots, or'if fibrous-rooted, then the lower cauline leaves narrow and less than 6.0 III wide; plants from western Helicon..."............................ 1" F. Biennial or short-lived perennieldfraa a short taproot or fibrous-rooted caudex, the stem-root Junction often con- spicuously thickened; stone erect or ascending, rarely de- cmbent; lower cauline leaves narrowly spathulate to ob- lanoeolate, less than 6.0 m wide; plants fro Chihuahua and Mango............................. 8. A. m 1". Annals frat a elenbr taproot, often Idth fibrous lateral roots; stems commonly decuabent, the central ones ascending; lower cauline leaves methulate or oblanoeolate to lanoeo- late, emetims subulate, cmonly more than 6.0 It wide; plants fro Jalisco and Riemann 9. A. condinentuu 0. W3 lavender, bright blue or purple wbn dry.................. 0 -95- G. Perennials from a fibrous-rooted caudexs basal leaves mostly persistent, with lug-tapering leaf bases; stems i‘ew, decmbent or ascending, simple or rarely asce- branched below the middle; midrib or cauline leaves not quite cartel-sung into the spices; multiple creme often present and the plants then netted................. H H. Achenes evenly glochidiate-pubescent; rays lavender throughout; plants from southern New been and eastern.I‘ameulipas................ 6. A. glendens H. Achenes glabrous, rarely sparsely glochidiate. pubescent nostly around the apex; ms lavender throughout or with alternating bands of light and dark lavender below, paler above; plants i‘rm Cerro Potosi, Nuevo Ieon................ 7. A. hmnii G. Annuals, fibrous- or taprooted; basal leaves soon deci- duous; stems erect, ascending, or decmbent, mostly (in nature moi-ens) nuch-brancbd below to above the middle; nidrib extending into the apiece-of tin leaves; multiple crowns never present...................... I I. Fibrous- or taprooted; stene canon]; erect or ascending; cauline leaves not densely crowded; plants from northern lluevo Don, and the south- central and east-central limited States.......... ............................ S. A. integglgolina I. wrooteds st-s nostly decubent, with densely crowbd leaves; plants tron the bane Pecos region 0: Me............u......... 10. Ae "bun“ 1. Astrsnthilan ylmodum Larsen Astranthim m Larsen, Ann. Mo. Bot. Gard. 20: 31. 1933. Keerlia mexlcana A. Gray ex 8. Wats., Proc. Am. Acad. 22: h22. 138?. hrpex JALISCO: Rio Blanco , 16 July 1886, 2.9.12.2! M (holotype, 0H2; isotype, Pm). Peremial, to 50.0 on high, from: a simple or branched, slender to stout woody taproot; stone 1 - several, erect or ascending, simple or loosely branched about the middle, greenish-brown to purplish, grooved and striate, essentially glabrous to densely strigose with coarse trichomes; peduncles 7.0 - ”.0 on long, noderately to densely white-pubescent below the heads; basal leaves not seen and not ob- served in greenhouse-grown seedlings; cauline leaves nearly glabrous to evenly and densely pubescent with coarse trichonss, up to 2.0 an long,nostly prominently three-nerved, the midrib conspicuous above and below, the margins entire, often revolute, sometime purplish, short- or long-ciliate, the spices obtuse or nucronate; lower cauline leaves reduced in size, 0.8 - 3.0 on long, 0.2 - 1.1 cm wide, spathulate to oblanoeolate, soon shriveling; middle cauline leaves oblong, ellip- tic, or oblanoeolate, 1.9 - 5.0 on long, 0.5 - 1.3 a wide, upper cauline leaves sinilar in shape and as large as the middle leaves or somewhat reduced, but rarely as sell as the lower cauline leaves; heads 1 - 8, rarely ncre, large, solitary on the pedmcles; receptacle low-conical; involucre cupanulate, 0.5 - 1.0 on high, 0.9 - 1.5 on -96- I’J CD Plate 9. Az-tmanthium niogpdmn, drawn from 93 Jon; 101 -98- n n,» .1“ , xx , ‘, Ni: 5 \\ I.“ «\V/ a. \NDII’I‘VWV I, .\ » lr .4 Kym/fled... xw {\“Kl‘D/yyy \ J / ‘1‘ v, «I» V 4%.. / 'I/fl , / 1 fl , 10h W . r -99- wide; pwllaries inbrioated in 2 or 3 series, the outer-oat ovate- lanceolate to linear-lanceolete, 6.0 - 8.5 II long, 1.0 - 2.1 me wide, nearly herbaceous throuhcut or with narrow to conspicuous scarious margins, these lacerate-ciliate free below the niddle to the apex, sparsely to densely pubescent on the back, glabrous and shim within, with often purplish, acute to acueinate spices; second series linear- lanceolate, 6.0 - 8.0 II long, 1.0 - 1.6 m wide, glabrous to moderately pubescent on the back; imenost (third) series, if present, 5.0 - 7.0 ewe long, 0.9 - 1.2 - wide, often Ivaline throughout; ray florets 15 - 30, white, sonetinee roseate when dried, the corolla 1.0 - 2.1 u long, 1.5 - 3.0 a- wide, the apex 2 - h-dentate or -denticulate; disk florets 90 - 120, the tube 3.1; - 13.1 nae long, slightly contracted at the throat, the lobes 0.9 - 1.0 as long; style branches of the disk with ovate or ovate-lanceolate appendages, about as long to one and a half tines as long as the stignatic lines; ray and disk achenes obovate, 2.5 - 3.2 - long, 1.1; - 1.7 II wide, with conspicuous ribs, the achenial faces nerveless or with 1 or 2 conspicuous nerves, glabrous, shiny, greenish-brown (vdth yellowish ribs and nerves) or dark-bran (with light-brown ribs and nerves), tin ray acmnes 2- or 3-ribbed (triquetreus when 3-ribbed), the disk achenes 2-ribbed or those nearest the rays often 3-ribbed and then triquctrous; pappus absent; chronosae umber 2 - 5, as detenined has no squashes. Plate 9. metribution: “stern Denice, at elevations between 800 and 1500 notes in tin Sierra liadre Occidental, in tin states of layerit, Jalisce, and llichcacan, in oak forests at lower elevations and pine- oak forests at higher elevatime. Principal flowering dates fru ~100- June to August. Plate 11. Qecinens examined: “1.13:0: pine-oak forest, Sierra del Halo, near lueber road to San Isidro, elev 11.90 a, 12 Aug 1961, pg £53 1339 (on; use; TEX; m; 05); in pine forest, Sierra del Halo, near lueber road to an Isidro, 010v 1530 a, n. m 1957, m 16101 (area; use); no memo , 16 July 1886, my; (holotype, 0m, isotype, PHI); shaded hillsides near Guadalajara, 27 June - 11; July 1893, m M}; (P; on; HIGH; HO; 14$; PH; 00). mimosa: oak forest, Coalcaeen, distr. Coalcaaan, elev 1250 n, 16 July 1939, m 1325: (on; 10; PH; 08). NAILRIT: oak forest along Bay from Topic to Coupostela, 7.5 mi n of Oaapostela, elev 800 n, 10 Aug 1961, p_s Aug 12110431), loan soil, wooded slopes and barrancas leading done) to lake NE of Santa Maria del Oro, elev ca 1000 m, 18 - 20 Aug 1959, W 621 (HIGH); pine forests innountains, ca 35 ni SWof Jaliscc, road to El Halinal, .1" 1150 - 1200 a, 19 June 1957, NM; (11108); red 01.: hills, nakforest sons, 7ailofcqcstela, e1ev800- 850a, 11Ju1y1957, “We“! 1.522 (HIGH). the prinitive characters of munit- m are discussed in greater detail in the section on the evolution of the genus. The species is unique in lstranthiu in its large, pruinently ribbed and oftennervedaohenes, leaves whicharehardlyreduced upward, and woody taproot. Astranthlu m requires relatively dry habitats in oak-, pine-oak-u or pine-forests at elevations which are the lowest for em species of Astranthiu. Its requir-ents of red clay soils ~101- parallel those of several other species in the genus. Astrauthiu m is sneewhat variable in pubescence. The type collection, m 1A6, and mpg, both from Jslieco, and the m 1222 specimens frat idchoscan are moderately to densely pubescent with coarse trichaaes up to 2.0 u long. a: the other hand, all other specisens exudned were essmtially glabrous or sparsely pubescent with trichones which are shorter and less coarse. The closest relationship of A. m is with A. laetifieu. Both specieshave'cln-aeosonenuebersmabase ofx-Sandtheir karyotypes show new sililarities. '33 echoes of A. lastificu marydnilartothcuofAMandthetwospeciesresenble one anotlnr in habit and leaf characters. Also, the distribution pattern of the two species, when correlated with norphcloglcal and cytological characters, indicates a close relationship. Letting.) lastificnn differs free A. am however, in its fibrous-rooted caudex, reduced pmllaries, abrqatly contracted disk florets, and tetraploid chruosme umber. ~102 - 2. Astranthiu laetificu De Jong, a). nov. Type: MICHOACAN: in Ling-Ages forest area, .Route 15, between Ens 2116 and 2&7, 1.5 ni W of fill Cmbres, elev 21190 n, 17 Aug 1961, 21,199; L199, (lblotype: H331; isotypes, am; 11:11; 00!; 08!). Perennis, osulibus simplicibus vel rsmosis, ascendentibus vel decmbentibus, ad 35.0 on ma, pubescentibus pills patentibus vel adpressis; folis basilaria spathulata, integra vel 1 - S-serrata; folia cauline integra vel l- S-serrsts, internedis spathulata ad oblanoeolate, 2.2 - 6.0 on longs, 0.5 - 1.5 on late, swericra pauci- reducta, suprua breve spathulsta ad plus ainusve lineal-is, 1.11 - 11.2 on low, 0.3 - 0.9 on late; peduculi circa 6.0 - 25.0 on longi, nudi vel cue brscteis solitariis, linearibus et minutis; oapitula solitaria, involucris 0.115 - 0.60 on altis, 0.9 - 1.3 on latis; plvllaria 2- vel 3-seriata, aequalis, late ovata ad ovato-lanceolata, spice acute vel longs scuinata, scarioso—narginata; flores radii circa 15 - 25, 1-seristi, albd; flores disci circa 80 - 110, corollis 3.3 - 3.7 In longis, flavis; aohsenia disci et radii cbovsta ad oblanoeolate, 2.5 - 3.5 a longs, 1.0 - 1.5 In late, empressa, glabra, sine pappo vel cue corona pappus linutissina; schaenia disci 2-costata, sed taaen achenia radii 2- vel 3-costata; chronosomaru 9. - 10. Perennial, to 35.0 on high, from a sinele or branching, vertical, oblique, or hcrisontal, fibrous-rooted cauden st-s l - several (rarely as law as 12), (hm-bent, sounding, or erect, simple or loosely branched below the aiddle, greenish to purple, sparsely to -103- - J ”W a n 1 -mh- ~105- noderately strigose; peduncles 6.0 - 25.0 cm long, naked.cr with a single, minute linear bract, the suits moderately to densely pubes- cent below the heads; basal leaves epsthulate, sparsely to moderately pubescent above and.below, becoming glabrous toward the long-tapering base, the margins short-ciliate and sonetines revolute, entire or scnetiaes with 1 - 5 callous-tipped teeth, the spices obtuse or aucrouate; cauline leaves sparsely pubescent above and.below, glabrous toward the base, the margins short-ciliate, entire, or with 1 - 5 shall, callous-tipped teeth, the spices aucrenete; lower cauline leaves shriyeling and.soon‘wanting, epsthulate, 1.h - h.5 cm.lcng, 0.3 - 1.2 ol.wide, the middle cauline leayes spathnlate to oblanoeolate, 2.2 - 6.0 on long, 0.5 - 1.5 cm wide, the mpermost short-spathulste to nearly linear, nostly reduced, 1.]; - h.2 on long, 0.3 - 0.9 on wide; receptacle conical; involucre c-panulate to heaispherical, 11.5 - 6.0 m high, 0.9 - 1.3 on wide; beds 1 - 8 (~12), solitary on the peduncles; phyllaries imbricated.in.2 series, rarely with a third, innenost series, about equal in length, 15.5 - 6.0 u long, 1.2 - 2.5 amlwide,‘broad1y ovate to ovate-lanceolate, the margins scarious, lacerate-ciliate above the middle, oftennpurplish, the greenemidp portion:deusely*pubescent on the back, the spices acute to long- sculinate; inner-est (third) series, if’present, linear-lanceolate, ‘withLverywuarrowumidpcrtiou.and.couspicucus scarious:margins or hyaline throughout; ray florets 15 - 25, white or>scmetmmes somewhat roseate ‘ neartb2-or3-tcotbdspea;diskflcrets80-110,thetube 3.3- 3.7 me long, strongly «attracted at the threat, the lobes ca. 1.0 u long; stamens with.subulste apical appendages and rounded anther bases; style branches of the disk‘with.narrowly ovate appendages -106- aboutaslongtoueeandahalftilesaslongastbstiuaticlines; achmes obovate to oblanoeolate, 2.5 - 3.5 .1 long, 1.0 - 1.5 - wide, glabrous, shiny, greenish-Iron to golden-brown, 2-ribbed or the ray achenes often 3-ribbed, the ribs of the sue color as tb achenial faces, the faces sonetimes with a superficial ridge; pappus absent or a minute whitish crown less than 0.1 3 high; chroncsue nuber g - 10, detemined free an squashes. Plate 10. Distribution; in the nountains of the 'lransvolcanic Belt in eastean Inchoacan, in pine-fir forest at ca. 2500 esters elevation. Mcwn only from the type locality and one additional locality near Morena. Flowering in August. Plate 11. ' Additional specimen emined; horelia, Jule-October, 181.0, Geleotti 2_l,2_o_ (P). the similarities between A. lastificu “A. m arcin- dicated in the discussion of tie latter. lh addition to being re- latedtoA.g_lgad_-_,A.laetificuisalsorelatedtcA.mssc_ege_ on the basis of the perennial habit, fibrous-rooted seeds: and open branchinghabit. m,the1eavescfA.Marencre nuercueandarelighterincclorandtexturethanthcsecfA. newt. Also, the heads of A. W are usually more nuerous and the achenes are 2-nsrved, never ribbed. 1h adlition, A.:_L£§_igo£isstetraplcidcnabasecf§-5whereasA.m misstetraplcidcnabaseofA-h. 1hroughA.lA§ifio-, itisposeihletorecogaisetherelationsMpsofA.muth theraainingspeciesongtrantMC. -107- Plate 11. Geographic distribution of A. m A. lastificne, A. pmurascens, and A. xanthoccmoides. ~108- 20 T 110 O A. XYLOPODUM 0 ALAETIFICUM l 110 I05 N w, NI 0 30 25 20 I10 IOS L I «it: IIO Q I'- 3. gtrsnthilsn gmusgens (Robins.) lersen “Rama“ 23223900118 (Rabi-nae) man, We MO. Bots Gard. 23: 33. 1933. Mammgm Robins” Proc. he. Acad. 27; 172. 1892. Iypes SAN LUIS POTOSI; shaded grassy slopes, Berranca of Les Canoes, 18 Aug 1891, ME (holctype, om; isotypes, Fl; 01-11; 1101, 2 sheets; MSCl; PHI, 2 shots; 00!; 08!). Perennial, to 60.0 cm high, from a short, vertical, airple or somewhat branched, fibrous-rooted cauder; stems l - numerous, sinple or repeatedly branched at or below the middle, erect or ascending, rarely decmbent, greenish to reddish-purple, glabrous to densely strigose with spreading hairs; pedmcles to 20.0 cm long, the emits sparsely to denseh white-pubescent below the heads; basal leaves spathulate to narrowly so, deciduous or persistent, essentially glabrous or marsely to moderately pubescent throughout above and below, or the upper son'face sparsely to moderately pubescent above themiddleandtowardtheapexandthonglabrous towardthebase, the lower sm'face sinilar to the wper or pubescent on the midrib only, the margins nearly glabrous to moderately ciliate-pubescent, the spices obtuse or nucronate; lower caulim leaves often shriveling, spathulete to lanceolate or oblanoeolate, 0.9 - 5.0 cm long, 0.2 - 1.8 cm wide, rarely longer and wider, the upper surface essentially glabrous to densely pubescent above the middle and then glabrous toward the base or glabrous to densely pubescent throughout, the lower surface glabrous -109- -110- Plate 12. Astranthiua pmurascfl, drawn from As. M 1221 (use). -112- or sparsely pubescent throughout, or Qarsely pubescent only above the middle and then glabrous toward the base concept for the pubescent midrib; middle and mper cauline similar to the lower in characters Of pubescence, the middle cauline leaves epsthulate, ovate, cbovate, or lanceolate to lance-elliptic, 0.7 - 5.8 on long, 0.2 - 1.7 cm wide, the spices minutely mucronate, chidate, or acuinate; upper cauline loaves lanceolate, lance-elliptic, or linear-lanceclate, 0.5 - 2.8 cm long, 1.0 - 8.8 an wide, the spices ouspidate to long-aclninste; receptacle conical, becaaing steeply so at maturity; heads 1 - numerous, small to large, solitu'y on the peduncles; involucre cnpanulate, 3.0 - 7.2 an high, 0.7 - 1.7 a wide; plwllaries imbricated in 2, rarely in 1 or 3, about equal series, broadly ovate to lanceolste, linear- lanceolate or oblanoeolate, 3.0 - 7.2 mm long, 0.9 - 2.1 an wide, with conspicuous to narrow, often pinkish to purplish scarious margins, these lacerate-oiliate mostly above the middle, the men midportions of the plwllaries glabrous to sparsely pubescent on the back, the spices ciliate-scnninate; ray florets 10 - 3), white, the lamina 0.9 - 1.1; cm long, 2.0 - 3.0 II wide, the spices 2- or 3-denticulate or -dentate, or conspicuously 2- or 3—c1eft; disk florets ho - 155, the tube cmtracted at the throat, 2.1 - 3.7 u long, the lobes 0.7 - 0.9 . 1mg; stylar appendages of the disk ovate toovate-lanceolate, half to about one and a half times the length of the etiolatic lines; acmnes obovate, glabrous to sparsely glochidiate-pubescent, canpressed or hardly so, 1.5 - 2.3 In long, 0.8 - 1.2 m wide, conspicuously or obscurely 2-nerved, greenish-breve: to reddish-brown, sometimes reddish- purple around the apex; pappus absent or a minute yellowish crown with a few teeth less than 0.1 m high; chromosaee nunber A - 8, determined -113- from Pm squashes. Plate 12. Distribution: in the flora Hadro m'iental, in tl'n states of San Luis l’otosi and mdalgo, and in the mountains of Oaxaca and Ghiapas, occurring prinoipally between 1700 and 2330 meters elevation. Habitats are open pastures, open pine forests, pine-osk-Juniper stands, oak forests, or oak-cypress woodland. Principal flowering dates tron June through November. Plate 11. anemone emined: 0W”: open pine-oak-Juniper stand, 9 mi NW of Omitan, elev 1790 n, 28 Aug 1961, am gag (GE; H803 TEX; 00; as); moist open pasture at San Nicolas, 5.2 mi SE of 'lbopisca, elev 1820 n, 25 Aug 1961, E 1295 1L6}; (GB; use; TEX; no; US); without locality or date, Ghiegbreggt 135 (033 P); without locality, 186).: - 1870, Ghiesbrem M (on; no); mountains along Route 190, about 5 mi E or San Cristobal do he Casas, ll June 1960, M182} (sion; m; as); wet fields, Just W of San Cristobal de las Cases, 21; June 1960, 92228 (HIGH; use; m; ac); pint-oak forest, about 10 mi E. o: utopia“, 2h June 1960, Q2}; (HEB; 11$; TEX; 00), sandy loan, pine-oak forests, in mountains along mute 190, about 9 mi W of Bonitan, 25 June 1960, m 3959 (HIGH; 14$; TEX; 00). W: nountain side, Jacala, elev 1500 ft, 25 June 1939. Lbs—0332 (I): roclq ravine, Jacala, elev 5000 ft. 5 My 1939. 253123 (1"; 083 HIGH: m); ate-p. N- teoing, pine-forested aloe at Village lilit of Cobrecito, between nuzaoanezm, sout- 85 olevzlma, 5 stpt 1961, 2mm (083 sec; m; as; E); mountainous area in mm Oak-m:- graced woodlands, 6.7 :1 s o: Jaoala, on noute 87, 28 July 1959. 355:3 -111“. (In). canals limestone slope along sout- 190, between his M2 and 1.13, elev 2130 m, 23 Aug 1961, a £332 (on: use; In; no; as); on grassy slope at tag. or oak forest, red clay loam, between nu 1.8!. and has, Route 190, 0.2 mi w of m hrtin, elev 2220 a, 30 Ana 1961, a m .l_1_1_6_(m13 11303 TEX; In; US). SAN LUIS P010813 shaded grassy slopes, Barranoa oi‘ Ins Ganoas, 18 Aug 1891, m1: £12 (holotype, OBI; isotypes, F83 am; 1401, 2 sbetss 1180!; Pitt, 2 sheets; m1; 081); without locality, 1891, £513}; gm 3.; (P); without locality, 1891, Virlet d'Agust m (P). n.- rame oi‘ Methim pmurascens is made q: or three popu- lations separated from one another by considerable distances. me northernmost or these populations is in the Sierra Hadre Mental in the states or San Luis Potosi and mdalgo. nu second population is inthmountains oanxacaandis separatedi‘ralthetirstbyadis- tanoe of acne £0 miles. The Oanoa population is in turn separated Iran tb third population in Ghiapas by a distance of about 250 miles. The plants or the three populations differ eqecially in charac- ters of pubescence, but these characters are too intangible to be or value in the separation of the plants. Thus. the Chiapas plants are essentially glabrous to sparsely pubescent and the achenes are neatly glabrous, rarely with a few glochidiate hairs. Contrariwise, the Oaxaca plants are sparsely to dousely pubescent and the achenes are mostly Qarsely glochiate-pubescent. m- San Luis Potosi and mdslgo plants are, on the whole, less pubescent than til Genoa plants but the achenes are glabrous or sparsely glochidiate-pubescent lib those ottheOhiapasplants. 'nlethreepwulationsare similar -115. in all other characters and their karyotypes are also similar. 01 this basis, it seems but to recoglise Astranthil- pmuragens as a single, if somewhat polymorphic Qeoies. Although the three populations are homoploid and belong to the sen! species, crosses between plants of the three populations yielded highly sterile Wide. lbtails on these crosses may be tomd in tin sectial a: croseim relationships in the genus. The relationships or gmthim pmmsoens are with A. laetgomu and with A. integi_:oliun. The similarities with tin latter were recognised by Larsen (1933). main integgoliun, however, is well set apart by the annual habit, smoother leaves, more numerous heads, and lavender rays. Their genetic kinship is indicated by the fact that both meoies hsve ohralosae numbers cal a base or g-h. A. pmurascens is also related to 5. ranthocanoddes. Although distinct from 5. pmurascens in its stolonii’erous habit, and con- siderably narrower leaves, 5. xanthooaeoides shares the one level or ploidy (g - 8), and both queoies have the soallest somatic chro- mosaue pair with satellites. h. Aetranthim xanthooanoides fleas.) lareen gtranthium manthocanoides floss.) Larsen, Ann. Ho. Bot. Gard. 20: 33. 1933. 31'me mthocaaoides less. am. Comp. 192. 1832. anemone xeranthocaloides Steud. Nmenol. But. 2: 220. 18140. (typographical error). £114 unthocomoides A. Gray ex Hensley, Biol. Centr. In. Bot. 23 118. 1881. Me: VERACRUZ: Prqle Ls an (In mya) , 29 June, M ggé (holotype, not known, possibly in HAL 1; isotype,. M01). Perennial, to 25.0 cm high, from a slender, fibrous-rooted caudex with mostly one erect flowering stem and few to many ascending or de- otanbent flowering stems and stolals; flowering stems 1 - many, simple or branched, green to purplish, sparsely to moderately strigose; stolons few to may, simple or branclnd, similar to the flowering stems in pubescence and ol‘ten bearing oapitula; basal leaves soon deciduous, narrowly oblanoeolate or gathulate, to h.0 cm long and on 1.0 - 1.7 cm wide, the blade nearly glabrous to moderately pubes- cent above and below, often glabrous toward the base, the margins ciliata, the apices mucronate; lower cauline leaves narrowly spathulate or oblanoeolate, glabrous to moderately pubescent above and below and then often toward the base, the margins ciliata, the spices pruninently mucronate; middle and upper leaves gradually reduced in size and similar to the lower cauline leaves in characters of pubescence, the uppermost -115- -117- Plate 13. Astranthiun xanthocomoides, drawn from 23 m 1125 (M30). -119- narrowly oblanoeolate to linear-lancoolate, the spices long-seminate; stolons basal, and then procmnbent, or cauline, up to no.0 cm long, with narrowly spathulate to linear-lanceolate, often minute (less than 5.0 m long) leaves, the tips of the stolons teminating in a capitulum or in a leery rosette, the latter taldng root and forming a new plant- 1et; peduncles to 12.0 cm long, naked, or with a minute, linear-lanceo- late bract; heads 1 - few, borne singly on stems or stolons; recep- tacle conical; involucre campanulate, h.2 - 7.2 mm high, 0.8 - 1.5 cm wide; pmllaries imbricated in two equal series, h.2 - 7.2 mm long, 1.0 - 1.8 mm wide, obovate to narrowly lanceolate, glabrous to sparsely pubescent on the back, the scarious margins lacerate-ciliate mostly above the middle, often purplish, the spices ciliata-sowinate; ray florets 15 - 26, white, sometimes roseate before anthosis and in drying, the lamina 0.9 - 1.h cm long, 2.1; - 3.0 mm wide, the apex entire or 2 - 3-toothed; disk florets 60 - 120, the limb contracted at the throat, 2.5 - 3.3 m long, the lobes 0.6 - 0.8 mm long; authors with minute subulate apical appendages; stylar appendages of the disk narrowly ovate, about as long to three quarters as long as the stig- matic lines; achenes obovate or narrowly ab, 1.14 - 2.0 mm long, 0.7 - 1.0 m wide, greenish-brown to dark-brown, nearly glabrous to mod- erately glochidiate-pubescent; pappus absent; oln'omosome nunber g - 8 dede iron Pm squashes. Plate 13. Distribution: in the mountains of mdalgo southtrard into the eastern mountains of the Transvoloanic Belt in Puebla, Tlaxcala, and Veracruz, principally at elevations between 2500 and 3000 meters. Habitats include moist, cool, open meadows and open pine- or fir -120- forests at high elevations and open pine-oak-Juniper forest at lower elevations. Principal flowering dates from April through October, with a single collection obtained in February. Plate 11. Specimens examined: HIDAIGO: ridge, ca 2 Ms S of Real. del Honte, elev ca 2880 m, 7 July 1959, m 2121 (use); without locality or date, Ehrenberg £1; (P), and Ehrenberg 6_1_6_ (P); open, grassy spots, between Pachuca and Real del Monte, elev 2700 m, 27 Aug 1911:, m 2.91 (F; on); open meadow in fir forest near Zereao and below Parque Nacional El Chico, elev 3100 m, 17 Oct 19146, m 1.529 (Gm; meadows of Sierra Pachuca, elev 9000 ft, 17 - 28 July 1898, m 9888 (F; GE; M0; NSC; PH; UC; US); alpine meadows, Sierra de Pachuca, elev 9500 ft, 26 Aug 1902, m 2828 (F; OH; HO); moist meadows and open woods, Pachuca, July 1905, M1213 (F; GE; M0; 00); Sierra de Pachuca, 21 6c 22 July 1901, £933 522; 93.1). PUEBLA: pine-oak-Juniper forest and green roadside, 5.3 mi Si or San Salvador El ace, elev ca 2500 m, 13 July 1960, .D_e_ £29; 62 (OH; M33; TEX; DC; US); on grassy slope at edge 01' pine tor-est, Route-119, 1.3 mi N or Tlaxcala state line, elev ca 2800 m, 2 Sept 1961, 23 m 12.9.6. (use); Cerro Azul, Feb 1909, Nicolas 21.9.: (F). mm. cool, moist, as... meadows, Route 119, 2.2 mi S of Puebla state line, elev 28h0 m, 2 Sept 1961, 95.9223 £91 (on; use; TEX; 00; as). mean: among dry turi' or in cultivated fields, Imogrande, lbunt Masha, elev 9700 ft, 27 April 1938, £14 331; (00; US); Joya, July 18hl, 'Liehaann 591 (P, 2 sheets); Props La Jays (La Bye) , 29 June, Schiede y; (holotype, not known, possibly in HAL ; isotype, H01). Larson (1933) indicated Iessing's name to be a "hunch subnudm'I but the brief description or the species is valid. leasing (1832) noted the collector as "ahiede", without locality or number, but in the emended description of the species by von azhlechtendal (1835), the locality and date are cited as I'l’r. 1a Jays Jun. 29". These data fit Schiede g_0_§_ in 140 which is considered to be an isotype. ggtranthim mthooomoides is a very distinctive species because or its conspicuous prooumbent stolons and reduced number of oapitula. Horphologioally, its closest relative is A. pu_rpurasoens with which it also agrees in chromosome nmnber and details of the karyotype. 'me distribution of the two species also indicates a close kinship. 5. Astranthium integgolimn (mom) Nutt. Astranthiun integggoliun (mchzd Nutt., Trans. Am. Phil. Soc. Ne Se 78 312a 1814]“ Key to the subspecies: Fibrous-rooted, sometimes indistinctly tap-rooted; achenes 1.6 - 2.2 mm long, 0.9 - 1.1 mm wide, glabrous or sometimes sparsely glochidiate-pubescent about the apex; involucre 11.0 - 6.0 m high, 1.0 - 1.5 cm wide; limb of the disk florets 2.0 - 3.2 mm long; ray corollas 1.2 - 1.5 cm long............... So. subsp. integgolim Tap-rooted, sometimes with fibrous secondary roots; achenes 1.0 - 1.6 mm long, 0.6 - 0.8 an wide, sparsely to densely and evenly glochidiate-pubescent; involucre 2.5 - 14.5 mm high, 0.5 - 1.2 cm wide; limb or the disk florets 1.5 - 2.3 mm long; rav corollas 0.65 - 1.2 a IMOOOOOOOOOIOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOSb. Bubape ciliata“ 5a. Astranthim intemum (Elohim) Nutt. subsp. integi_i‘olimn. gig; mugglia mean, 21. Ber. at. 2. 131. 1803. Type; WESSEE: "lh mense maio floret Gtmberland', 1795. m 2.1.9.... (holotype: P , photo, arm, 01‘. discussion in the historical account or gtranthi ). ' Lngmm" New 11. u. an. 2. 23. 1837. Type: unknom. Inoality: "Kentuclq". wagging” New 11. w. an. 2: 23. 1837. mm unknown. -122- -12 3- locality: "test Kentuclq'. £1133 integiiolialiiohx. var. M Raf., New F1. N. In 2. 2h. 1837. Type: unknown. locality: unknom. 3121114 integgglolia Hichx. var. _e_I_1._a_t_gRsf., New 1‘1. N. Am. 2: 214. 1837. Type: unknown. locality: mknown. I 2111; intemgolia Michm var. m Bath, New Fl. N. Am. 2: 21:. 1837. Type: unknown. locality: tmlmown. Annual, to 1:5 .0 cm high, fibrous-rooted or smetimss tap-rooted but then with amorous secondary roots; stone 1 - ntsnerous, erect, ascending, or sanetimes spreading, simple or loosely branched iron below to well above the middle, essentially glabrous to moderately strigose with fine, appressed and spreading trichomes, often densely pubescent toward the base, greenish to reddish-purple; peduncles to 20.0 cm long, sparsely to densely pubescent below the heads; basal leaves narrowly to broadly epsthulate, 1.8 - 8.0 cm long, 0.6 - 2.2 cm wide, with long-tapering bases, the lamina Qarsely pubescent above with fine trichases, especially on the mper half and then glabrous toward tin base, glabrous below or ally with a few scattered hairs on the prominent midrib, the margins finely ciliate, the spices obtuse or mucronate; cauline leaves reduced upward, sparsely pubescent above, especially toward the apex and then glabrous toward the base, glabrous below or with scattered hairs on tln midrib; lower cauline leaves spathulate, oblanoeolate, or lanceolate, 2.3 - 5.8 cm long, 0.6 - 1.7 cm wide, the margins ciliate, tin spices mucronate; middle cauline leaves elliptic to lanceolate or oblanoeolate, 1.5 - 5.5 (-6.7)cm long, 0.3 - 1.5 {-2.1} cm wide, the midrib sparsely to -12),- moderately pubescent below, the margins and spices similar to those of the lower cauline leaves; upper cauline leaves oblanoeolate to lanceo- late or linear-lanceolate, 0.8 - 3.6 cm long, 0.1 - 1.2 cm wide, some- times glabrous throughout, the margins ciliate, the spices acute or acuminate; heads 1 - numerous, borne singly on the peduncles; receptacle conical; involucre campsnulate to hemispherical, 1:.0 - 6.0 mm high, 1.0 - 1.5 cm wide; phyllaries imbricated in two equal series or in a single series in small, secondary heads, 1:.0 - 6.0 nm long, 1.0 - 2.0 m wide, ovate or ovate-lanceolate to linear-lanceolate, the margins scarious, lacerate or lacerate-oiliate mostly above the middle, the green midportion glabrous or with a few scattered trichomes on the back, the spices acuminate to long-acminate; ray florets 114 - 3), white or more commonly lavender below and pale above, fading in m, drying purplish-blue or dark-blue , eqecially on the distal half, the corolla 1.2 - 1.5 cm long, 3.0 - h.6 mm wide, the apex entire or 2- or 3—toothed; disk florets 100 - 210, the limb 2.0 - 3.2 mm long, abruptly contracted at the throat, the lobes ca 0.5 m long; stylar appendages of the disk ovate, about as long to one and a half times as long as the stigmatio lines; achenes obovate, 1.6 -.2.2 on long, 0.9 - 1.1 mm wide, glabrous or sparsely glochidiate-pubescent mostly about the apex, often within the same capitulun, greenish-brown or dark brown; pappus absent or a mall, whitish or yellowish crown less than 0.1 m high; chronosone nunber g - 1:, determined from Pic squashes. matribution: 1mm central Kentuclq south to central Tennessee, extreme northwestern Georgia and northeastern Alabama, west into Missouri and Arkansas where it hybridises with subsp. ciliatum. A -125- single collection has been obtained in central Mississippi. Principal flowering dates iron May through July. Plate 11.. ' Representative specimens: ALABAMA: JACKSON 00.: rocky banks in the mountains, aottsboro, 2 May 1889, in; 1,; (use). ARKANSAS: HOT arms 00.: not airings, May 1879, m L2; (no 122885). 14153315an 00.: Joiner, 11 June 1931, gy_l_e_ 662 (TEX). wlssmnm 00.: Fayetteville, 1880, 3521 M. (MO 122880). GEORGIA: DADE 00.: dry limestone ridges near Trenton, 20 May 1899, Biltmore Ibrbariua N9 £6082 (CH); dry grounds, Ringold Road, Chickamauga, 27 May 1911, Churchill 122 (CH; 110, 2 sheets; TENN). KENTIBKI: ADAIR 00.: alluvial field, near Green River, 10 mi N of Columbia, 1 May 1959, 91-9329 8: m 21 (P). All“ 00.: road- side at edge of woods, in dry ground, Adolphus, 21 my 1933, thatherbz _& Weathers): 622 (CH; TENN). EDMONDSON 00.: vicinity of Namath Cave, May 1899, 25139333:- (GH). ESTILL C0.: Irvine, June 1831:, £33.33; £1.21 (HIGH). MADISON C0.: fields and roadsides, between Big H.121. and Kingston, 9 Hay 1939. max; (F): 812 8111, and the valley be- low it, in meadows, 1 June 1833, mg; (PH); bank of creek, 1.75 mi NE of Big Bill, 17 key 1911;, M9291 (rm). 14mm: 00.: lowg'roundnear Buckhom Creek, 6miSHoflebanon, 91101931, mm (m). Wm 00.: damp soil, wooded flats of the nicks River, Burgin, 10 June 1916, Egg (F). 1131501 00.: limestone soil, edge of thickets, Balltown, 18 may 191:0, m 321; (F); in woods, at. Thomas, 10 Hay 1911:, mum 32 (HIGH; as), dry roadside near Balltown, 22 Hay 1933, leathery é Weatherbz 622 (GR). 311881311. 00.: along roadside under oak-guI-sourwood, about 5 mi N of Russell -126- alrings, June 1961, m E} (F). WARREN 00.: Barren fiver, banks, May 1890, gr_i_o_e_ £1.13; (drawing, 140122883): Bowling Green, 6 May 1892, £133 M; (MO 122882) 3 river bluffs, Bowling Green, May 1896, gr_i_o_e_ _s_._g_1_.. (MICH) . WAYNE 00.: low grassy bank of Beaver Greek, 3:! of Monticello, 12 - 11: July 1937, mg m 21; (F; 011). MISSISSIPPI: HOLMES 00.: 'l‘cbula, 18 April 1927 , M if. Anderson 1252 (M0). TMESSEE: CHEATHAM 00.: wooded cove of bluffs, Harpeth River, along Route 70 3, ca 18 mi 3:! of Nashville, 8 May 1958, M321; (SW): low woods, Pegram, 6 May 1939, MM (01!; UC). DAVIDSON CO.: calcareous bluffs of Ehrpeth River, ca 15 mi sw of Nashville, 11 my 1958, Shannon 1231 (am); linestone slope w of Nashville, 17 May 1931:, m 1813 (OH; mm); open cedar woods, without locality, 3 June 1923, M 33.49. ( PH): moist field, Joelton, 16 July 1922, m l_l§_ (on). 00011012 00.: without locality, May 1853, 312.2% (HIGH). HICKMAN 00.: moist, readside ravine, between Graham and nunnelly, 20 April 1909, M 23 3;. 111322 (Tm). HOUSTON 00.: meadow near Wells Creek, Erin, 18 my 193:, m 182 (011: mm). uoumam 00.: Dmlbar Gave, 25 April 1912, m 182,: (Tim): omningban Lake, 8 mi an of Clarksville, 1 May 1908, Olebsch £112 (mm). RUTHERFORD 00.: off Us Routes 1:1 and 70 3, ca 1: mi N of Lavergne, 18 may 1958, Channell 121,; (mu). 31mm 00.: near Dover, 8 May 191:2, 99931263 (TENN). wnsou 00.: rocky glade, Cedar of lebanon State Park, 10 mi 3 of Lebanon, 11 May 191:1, m 9325 (310). mm 00.: open woods and roadside, near Sparta, 13 May 1933, Neatherbz g Weatheryy 6_231 (on) . 00mm mm: "In mouse maio floret Cmnberland", Michaux 2.2.2:. (holotype: P , photo 1432!). -127- Plate 11:. Geographic distribution of Astranthium integgolium subsp. integrifolium, _A_. integifoliun subsp. ciliatum, and A. robustun. -128- zamamomd .. i I a e 220252.: N: 22.65532. .23... 2:38.5334 eo , ¥ 22.5.5 .32.. 22.non_¢ouhz..< Kneee .e A e e e e e .v x . o . l t.” .1. . “a e e tawny: a. . ' “.. ‘«1‘ C 1 d w A e a e e e e e a h o e e ee e O l O C e v 0 O . e ee e e afiee ee ‘0 . o e e e OJ .u . . e e e O o . C. . O %% . e ee ‘ ‘ r9 e . . o u 00 o 0 OO O O -129- Sb. Astranthium inteegolium (Michx.) Nutt. subsp. ciliatun (BIL) m Jong, stat. nov. giggle—3333f" New Fl. N. Am. 2: 21:. 1837. 17130.8 TEXAS: "San Felipe de Austin", 1835, Ian-amend 2;; (leototype, 1m, photo, M8013 isotypes, 0m; GL1: K 3 N11). 21133311333“. var. triflora Raf., New Fl. N. Am. 2: 25. 1837. Type: unlmown. Locality: ”Pens”. Astranthium integi_foli\an (310111.) Nutt. var. ciliatun (Rah) Larsen, Ann. 1%. Bot. Gard. 20: 31. 1933. Astrantlrun inteflolium ( )flohz.) Nutt. var. rosulattan Larsen, Ann. 140. Bot. Gard. 20: fl. 1933. Me: TEXAS: Matagorda 00.: on sandy prairies, Matagorda, 5 Hal-ch 1911:,_l;a_]_;_m_e_r_ i852 (holctype, H01). Annual, to 33.0 cm high, with a taproot or smetimes with fibrous secondary roots: stems 1 - nunberous, erect or ascending, simple or loosely branched fro: below to above the middle, glabrous to sparsely strigose, often densely strigose near the base, greenish to reddish- purple: peduncles to 10.0 on long, sparsely to densely pubescent below the Dads: basal leaves narrowly spathulate, 1.2 - 5.1: cm long, 0.5 - 1.3 cm wide, sparsely pubescent above, emacially toward the apex and then glabrous toward the base, glabrous below or with a few scattered hairs, the margins ciliate, the spices obtuse or mucronate; cauline leaves reduced upward, sparsely to moderately pubescent above or only so toward the distal end and then glabrous toward the base, glabrous or warsely to moderately pubescent on the midrib below, the -13).. margins ciliate: lower cauline leaves narrowly wathulate to oblanoeo- late, l.5 - h.O cm long, 0.1: - 1.2 cs: wide, the margins often puaplieh, the spices mucronate; middle cauline leaves similar in shape, 0.6 - 2.7 cm long, 1.6 - 7.0 m wide, the spices mucronate; upper cauline leaves narrowly lanceolate to linear-lanceolate, 0.6 - 2.1 cm long, 1.0 - 1:.0 m wide, the spices acminate: bade l - numerous, borne singly on the peduncles: receptacle conical: involucre campanulate to hemisphe- rical, 2.5 - 11.5 m high, 0.5 - 1.2 cm wide; p1wllaries imbrioated in two equal series, ovate, lance-elliptic, or ovate-lanceolate to linear- lanceolate, 2.5 - 1:.5 m long, 0.8 - 1.8 m wide, the margins scarious, prominently lacerate-ciliate above the middle, the green midportion glabrous or sparsely pubescent on the back, the spices ciliate-aotaain- ate : ray florets 1h - 35, white, or lavender below and paler above especially towud the distal end, 0.65 - 1.2 cm long, 2.0 - 3.1 mm wide, the apex entire or 2- or 3-toothed: disk florets 120 - 170, the limb 1.5 - 2.3 m long, abruptly contracted at the throat, the lobes 0.5 - 0.6 m long; stylar appendages of the disk ovate, about as long to twice as long as t1. stignatic lines: achenes obovate, sometimes narrowly so, l.O - 1.6 u long, 0.6 - 0.8 an wide, marsely to densely and evenly glochidiate-pubescent, the apical hairs smetimes so dense as to simulate a pappus crown, rarely a few acbnes glabrous and then mostly within the one oapitula, greedah-brom to dark-brown: pappus absent or a minute, whitish or yellowish crown less thm 0.1 an high: chrmosone nmber g - 1:, determined fro: Pm equashes. Distribution: frua suave neon, lance, northward into east- central and eastern lens, eastern Oldnhona, and southermost Kansas, -131- east into Missouri and Arkansas where it lubridises with subsp. integgiioliun. This widegread taxon occupies a wide array of habi- tats, ranflng from dry to moist open pastures, roadsides, and native prairie to oak-pine , oak-elm, oak-hickory, and oak-pine-hicloory wood- lands. Data from barbarium specimens indicate a wide range of soil types, fran sandy soils to saw-loam, saw-clay, and clay soils. Principal flowering dates in April ad m, with a few early collections obtained in March, and a few late ones in June and July. Plate 11:. Representative specimens: ARKANSAS: CIEBIIRNE 00.: dry bluffs, Van Bln'en, elev 550 ft, 29 June 1937, m 2:522 (P: M0); dry rocky ridges, Tunbling Shoals, elev 1:50 ft, 19 no 1951, mm (am, 2 sheets). cameos 00.: Pine Swings, 8 Hay 1938, mm (um). um 00.: Cedar Park, near Coma, 17 M 1933, Egg 2:9; (mm). HOT SPEEDS 00.: rocky creek bottoms, Chamberlain Creek, Hamlet Cove, elev 550 ft, 2 new 1937, Demaree 11:92; (I; no): Chamberlain Creek bottoms, Magnet Cove, elev 1:90 ft, 1 my 1938, m 1.7321510”; Chamberlain Creek bottans, Magnet Cove, elev 1:50 It, 30 April 1939, nemeree 1&2. (no: 340: 00). W 00.: call roclq creek bottoms, Saline fiver, Dierks, elev 1:20 ft, 3 no 1955. Deneree m (we: aw). Jamaal 00.: glade, lmiNHofPi-W: Burma, mutgmmo). W 00.: rocky wooded creek bottaa, Cadio Gap, 23 say 1951:, Demaree 529. (am, 2 shots). POLK 00.: bottus of Cassetot mver, elev 1000 It, 3 an 1955. mg: (mm; 3:8). mm 00.: Little Book, 10 Hay 1885, £03 g_._n_,_ (P). SHINE 00.: lull creek bank, branch of Alan Pork, state hospital, elev 260 ft, 28 may 1951:, 432- ' mm (mm: 340, 2 sheets). SEBASTIAN 00.: s of Greenwood, 2 Maw 1951, m 292.“; (so). W 00.: roclq creek bottoms, St. Joe, elev 700 ft, 21 June 191:1, m 7.12393 (no: 8148). Kansas: 0310110001: 00.: rocky soil, 7 New 1897, M 1955 (on: no): mall, overgraaed pasture in scrub-oak woodland, sandy soil, 3 mi E of Sedan, 2 W 1961, mango; 1% (RAND: n30). OWL: ADAIR 00.: along bank of little Dee Creek, 15 mi ass of Stillwell, 6 May 1950, mtOMnson 1:2 (010.): in oak-hickory woods, hill near Brush Mountains, 12mi Sof Stillwell, onusS9, 12m1958, 1%6_§_1_2_(mu: elm; TEX). ATOKA 00.: 7 mi w of Atoka, clay soil, woods and ravine near as. 19 my 19th. miss; merino-116.90% MW M sandy clay, roadside, I: mi s of Kiowa, 28 April 1955, sxinnere 1218; (340). 011mm 00.: wet woods and 0113, Devil's Owen. 5 mi 0 of Hinton, 25 April 1937, g 1391 (on). 0mm 00.: loan soil, Hurray Lab, 11 April 1939, mg (0m): clay washes near Ardaore, 17 April 1913, _s_tg_ven_g 31(03). CHEROEE 00.: rocky sanw loam soil, lepadden Falls, 7.8 mi us of Tableqush, 2 my 191:7, m 29, (TEX): prairie area, 3miEof Ft. (atom, on state 10 andlmiN, 26 April 1958, wanie M11. (mu; em: all). CLEVELAND 00.: dry, olayiehupland,l:adao:wo:aan, zhmu11928,;a_a_r__klez£_._n_._ (0mm: prairies, sorremton, 6m1931, gage-egg. (on. 1.23.111): 00:1. 00.: dry, rocky hilleide, Topelo, 10 april 1936, De-aree 1.1212(0nh 004010118 00.: sent loam soil, meadow, 13:]. E of Lawton, lei s, 26m19ln,gg_gh_1_(on.):eandyloueoil, lbdioine Park, 17 April 1937, 23311 ( OKLA): moist soil, inchita maintains, 7 Kay 1921, $.29; 231 (on). CMTAW 00.: sanu loam soil, -133- sniaoraugo, vmwhmngmm. 03118100.: Sapulpa, 19 June 1891:, 213322; (on: no): gravelly clay hillside, 13.5ai a of Mt, 8 June 1928, m 1.2 (OKLA): waste places, Bristow, 3w19ho, main: 2 (on). momma meadow soil, 3ndwo: Keota, 29 April 1939, _0_1_ar__15 a m g_2_ (on). amass 00.: dry pasture along Hwy 12, 1:.7 mi E of county line between Hughes and Pontotoc counties, 20 April 1961, E. 933182 (1930); dry roadside, 6 mi E of Holdenville, 7 May 1937, my; (010.). marten: 00.: vauriloa Lake, near wauriloa, 15 April 1957, mg 2.2.2.: (on). 00313101: 00.: moist, unshaded creeldoank near Hamsville, April 1916, Griffith 3:55 (GK): open woods and rocky glade in Tishaaingo granite fomation, 6 mi 8 of Hill Creek, 29 Hay 191:0, m 122 (on): in weeds and meadows, near Tishomingo, 27 April 1916, 8)th 33,: (GH). m 00.: Toulouse, 11 Hay 1907, M28 (P): sandy clay soil, E of clay Inna, maohven, 29 April 191:5. EL”. 2: (OKLA). LATHE! 00.: inrichneadow along noeklelendn. ante: Highs, 21mm, nulls; E52: emawms mob-a W: 9W 1937, Standish 23 (01:11). W 00.: Goat's Bluff, near Smerfield, elem stress, 27 April 19%, 2.9.9:. at: (on, 2 sheets): upland prairies, saw soil, near Howe, 25 m 1931, Palmer 23} (on; no). LOVE 00.: 8W 10“: Lab Wrapark, 21: April 1937: Beard ,3}, (010-)- mm 00.: rocky soil in wood, near Cup $0tt, locust Grow, 3 w 191:0, mnmm): prairie area, halfanile NandleiWofPeggs on State 82, 11 w 1958, 3133 6315 (mu, sun; in). woman: 00.: oak-pinewoods, Beaver: BendState Park, 6miN, hmiEof BrokenBow, 18 mull 1959, gig-gm (RAND): open woods, near Idabel, 18 Hay 1916, m 395 (cm: oak-biolmry-pine forest along road between the 431,- Narrows and Smithville, 8 June 1930, 9219, gag-103111; (0K1). HCINTOSH 00.: prairies along railroad, Checotah Road, 2 Hay 1939, flfig (on). W 00.: Arbuckle Homtains, Crusher, 12 Hay 1916, Enig 606 (MO): moist soil, banks of Washita, near Dougherty, 25 Hay 1907, Rutledge g 953, g_.__n, (on. 12139): SE of Sulphur, 13 April 1926, .3229. g5 (on). HUSMEE 00.: prairie, roadside, moist black loam soil, 1 mi NE of Muskogee, 9 May 1926, my ( on); sand-waste field, 3 mi E of Bragg, April 1950, m 6_2_ (SID): floodplain of little Greenleaf Creek, E of cup oruber, 6 say 1950, Dohrbaggh 11; (OICL). NOBLE 00.: sandy 01v soil, 2.5 mi E of Morrison, 1 m 1937, m _s_.__l;._ (on; am 20162). 01041101: 00.: moist soil, pasture, 3111 130de city, llay192h, mama); aandy soilin open postoak-blackjack woods, 6 mi E of Britten, 17 14v 191:1, Waterfall 2_70_2_ (on; on). 0mm 00.: sandy clay loan, 8 mi 3, 2 mi 3 of Mounds, 25 March 1937, m 39 (0114:). cam 00.: clay loam soil, 6 an. NE of Prue, 21: April 1937, 0333-31 L0 (OKL): saw soil, oak-hickory open wooded area, 8 mi N of Pawhuslca, 3 May 1958, 1::ng mg (mu): Boy Scout Cup, 15 mi w of Bartlesville, 10 Hay 191:1, §_i_c_e_ Q; (on). MINE 00.: low spot in pasture, 6 mi N of Stillwater, 15 June 1951, 93121; 222 (OKLA): loam soil, near Stillwater, l m 1935, m 11 (PH: TEX). PITTSBUK} 00.: open woods, E of Hartshorne, 11 June 19”, _Ol_a_r_lg {£22 (on): dry pasture elem Hey 270, 6.5 mi 8 or Arpellar, 20 April 1961, p_e_ m 181 (um): roch soil, open blackjack forest, Dab lelester, 15 bril 191:1, may, (0m): black sandy loam, 3.1: mi N of Kiowa, 28 mril 1955, mm (340). PGITOTOO 00.: lo- sodl, 1: mi 8 of Stonewall, 16 April 19$, m 125 (TEX): well-drained meadow in vanteraith Park, 1135.. 1 mi SE of Ada, 2 April 1951, @123 (om); oak-hickory open woods, 5 mi 3 or Allen, 10 Any 1952, 521219; (on). PUMAHMIE 00.: woodland, 1 mi SW 01' bomseh, 28 June 1937, 13515131 1:222 (H03 on); prairie, w o: Pink, 3 April 1932, mg ( on). 2mm 00.: roadside, 8 mi NE of Antlers, 16 April 1951;, Mg}; (031.); 1 mi s or Nashoba, 31 Hay 1919, gags 3g; (in): am rock: oak-elm woods, 3 mi w or Corinne, 15 April 1955, mg: (03; HIGH): dry, roclor barrens,hmiEotA1bdon, 9&91937,§gpy£am118_8_(om; hmi NEot Finlay, 9 no: 1937, mg (on). man co.: open fields and roadsifis near “awoke, 12 Hey 19h0, M 62 (on). SEQUOIAH 00.: open field and roadside at Mam Mission, 6 mi NE of Sallisaw, 7 m 1955, @2291; (cm); oak woods at mull» Laloe, 5.5miNEo£00re, onstuto 100, 7mI957, _th_i_._g£1_§(om). moms co.: catoou, 8 In 1895, E3 821 (m). swarms co.: 3 mi 31 or Marlow, 2 Hey 19141, 92 E. (on). 201.31 00.: may 1on. road- side, 3 oi Sand airings, 30 April 1938, Baldook g; (m); oak-hickory association, vicinity of Tulsa, 29 April 1937, Luokharot a: (on). wmm 00.: loan of prairie, W, 28 an 1920, 1.9—“ .129; (PH). TEXAS: W00“ oak-pinewoodemgentle Mes, InhView intnodiot Asa-n1; aromas, 12 April 1955, mm (on; m); near Palestine, 1 mil 1935, whitohouoo _1o_2;6_2_ (ma; an). Annals 00.: Aromas usage, 9 April 1985, 995198259 (on; an). Ami-rm no.3 dry sandy soil, SenhlipedemeflnStatePark, suitors-my, 28 Haroh19h9, mm